Dermal Shield Research Articles

A new old Budgett frog: an articulated skeleton of an Early Pliocene Lepidobatrachus (Anura, Ceratophryidae) from western Argentina

ABSTRACT The South American horned frogs or ‘escuerzos’ (Ceratophryidae) are among the best-known neobatrachians and comprise the genera Chacophrys, Ceratophrys, and Lepidobatrachus. Their relatively rich fossil record, which dates back to the Late Miocene and is largely concentrated in the Argentinean Pampas, consists mostly of variably complete cranial remains and comparatively few, mainly isolated, postcranial bones of Ceratophrys, whereas only three specimens of Lepidobatrachus are known. Here we describe two extra-Pampean ceratophryid records from the Upper Miocene–Lower Pliocene of Huayquerías del Este, Mendoza Province, Argentina, including Lepidobatrachus dibumartinez sp. nov. based on a well-preserved articulated skeleton. The new species is part of the stem of Lepidobatrachus and is diagnosed on the basis of several cranial and postcranial traits, including a large kite-shaped dorsal dermal shield. The new taxon sheds light on the evolution of a dorsal shield in the group, as well as on their paleobiogeographic history, adding support to the hypothesis of a broader ancestral distribution of Lepidobatrachus, whose diversification might have been triggered by the Late Miocene marine introgression of the Paranaense Sea. Also, it indicates that the prevalent climatic conditions during the Early Pliocene in western Argentina were much warmer and less dry than today. Finally, we discuss the potential value of the new records in calibrating the time-tree of Ceratophryidae. http://zoobank.org/urn:lsid:zoobank.org:pub:2DFB3B68-8C30-4252-A3DC-BF1F69F5AB6A

The evolution of dermal shield vascularization in Testudinata and Pseudosuchia: phylogenetic constraints versus ecophysiological adaptations.

Studies on living turtles have demonstrated that shells are involved in the resistance to hypoxia during apnea via bone acidosis buffering; a process which is complemented with cutaneous respiration, transpharyngeal and cloacal gas exchanges in the soft-shell turtles. Bone acidosis buffering during apnea has also been identified in crocodylian osteoderms, which are also known to employ heat transfer when basking. Although diverse, many of these functions rely on one common trait: the vascularization of the dermal shield. Here, we test whether the above ecophysiological functions played an adaptive role in the evolutionary transitions between land and aquatic environments in both Pseudosuchia and Testudinata. To do so, we measured the bone porosity as a proxy for vascular density in a set of dermal plates before performing phylogenetic comparative analyses. For both lineages, the dermal plate porosity obviously varies depending on the animal lifestyle, but these variations prove to be highly driven by phylogenetic relationships. We argue that the complexity of multi-functional roles of the post-cranial dermal skeleton in both Pseudosuchia and Testudinata probably is the reason for a lack of obvious physiological signal, and we discuss the role of the dermal shield vascularization in the evolution of these groups.This article is part of the theme issue ‘Vertebrate palaeophysiology’.

Description and phylogenetic relationships of a large-bodied sphagesaurid notosuchian from the Upper Cretaceous Adamantina Formation, Bauru Group, São Paulo, southeastern Brazil

In this work we describe new remains and possible gastroliths of a sphagesaurid (Mesoeucrocodylia: Notosuchia) unearthed from the Adamantina Formation (Bauru Group, Upper Cretaceous) in the municipality of Fernandópolis-SP, which add new data about the dental, dermal shield, and the post-cranial skeleton morphology of these crocodilians. Phylogenetic analyses place the studied fossil within Sphagesauridae, in a polytomy with Armadillosuchus arrudai and Caryonosuchus pricei. The anatomical comparisons are congruent to the phylogenetic analysis since they also suggest that the specimen herein described is closely related to other larger-bodied Sphagesauridae species, such as Armadillosuchus arrudai. The new skull, appendicular, and dermal elements described in this work may provide data for the elaboration of new phylogenetic characters which can improve cladistic analyses concerning Sphagesauridae species, as well as they may help to better understand the morphological complexity of this family of crocodylomorphs, especially the large-bodied species.

Redescription of Phymolepis cuifengshanensis (Antiarcha: Yunnanolepididae) using high-resolution computed tomography and new insights into anatomical details of the endocranium in antiarchs.

BackgroundYunnanolepidoids constitute either the most basal consecutive segments or the most primitive clade of antiarchs, a highly diversified jawed vertebrate group from the Silurian and Early Devonian periods. Although the general morphology of yunnanolepidoids is well established, their endocranial features remain largely unclear, thus hindering our further understanding of antiarch evolution, and early gnathostome evolution. Phymolepis cuifengshanensis, a yunnanolepidoid from the Early Devonian of southwestern China, is re-described in detail to reveal the information on endocranial anatomy and additional morphological data of head and trunk shields.MethodsWe scanned the material of P. cuifengshanensis using high-resolution computed tomography and generated virtual restorations to show the internal morphology of its dermal shield. The dorsal aspect of endocranium in P. cuifengshanensis was therefore inferred. The phylogenetic analysis of antiarchs was conducted based on a revised and expanded dataset that incorporates 10 new cranial characters.ResultsThe lateroventral fossa of trunk shield and Chang’s apparatus are three-dimensionally restored in P. cuifengshanensis. The canal that is positioned just anterior to the internal cavity of Chang’s apparatus probably corresponds to the rostrocaudal canal of euantiarchs. The endocranial morphology of P. cuifengshanensis corroborates a general pattern for yunnanolepidoids with additional characters distinguishing them from sinolepids and euantiarchs, such as a developed cranio-spinal process, an elongated endolymphatic duct, and a long occipital portion.DiscussionIn light of new data from Phymolepis and Yunnanolepis, we summarized the morphology on the visceral surface of head shield in antiarchs, and formulated an additional 10 characters for the phylogenetic analysis. These cranial characters exhibit a high degree of morphological disparity between major subgroups of antiarchs, and highlight the endocranial character evolution in antiarchs.

Do the ornamented osteoderms influence the heat conduction through the skin? A finite element analysis in Crocodylomorpha

In order to assess the implication of the crocodylomorph ornamented osteoderms on the skin conduction during basking, we have performed three dimensional modeling and finite element analyses on a sample which includes both extant dry bones and well-preserved fossils tracing back to the Early Jurassic. In purpose to reveal the possible implication of the superficial ornamentation on the osteoderm heat conduction, we repeated the simulation on an equivalent set of smoothed 3D-modeled osteoderms. The comparison of the results evidenced that the presence of the apical sculpture has no significant impact on the osteoderm global conduction. Furthermore, as we also aimed to assess the influence of the inner bone porosity on the osteoderm conduction, we modified the heat equation parameters so that the 3D-modeled osteoderms successively score the compact and the cancellous bone properties (i.e. mass density, heat capacity, thermal conductivity and thermal diffusivity). Finally, we repeated the analyses using the soft-dermis properties which lead to outline that neither the degree of porosity nor the presence of the osteoderms (in itself) significantly modifies the heat conduction through the crocodylomorph skin. Consequently, as hypothesized by previous authors, if the dermal shield happens to be involved into heat capture during basking for crocodylians, this process must mainly rely on a convective effect based on the osteoderm relative degree of vascularization. This last assumption could thus explain why the crocodylians which produce little metabolic heat would carry an entire vascularized osteoderm shield.

Osteoderms of Montealtosuchus arrudacamposi (Crocodyliformes, Peirosauridae) from the Turonian-Santonian (Upper Cretaceous) of Bauru Basin, Brazil

The skin of Crocodyliformes is characterized by osteoderms of various sizes and shapes. It plays roles in defence, thermoregulation, sexual attraction, calcium reserves, and locomotion. This study presents the morphometric characteristics of osteoderms preserved in the nuchal, dorsal, ventral and appendicular shield of Montealtosuchus arrudacamposi, a Peirosauridae crocodyliform from the Turonian-Santonian (Upper Cretaceous) of Adamantina Formation, Bauru Basin, Brazil. The results of the analysis show that its dermal shield had protective and thermoregulatory functions and a well-defined mechanical function allowing terrestrial locomotion and enabling good agility and capacity for movement.

Origins of bone repair in the armour of fossil fish: response to a deep wound by cells depositing dentine instead of dermal bone

The outer armour of fossil jawless fishes (Heterostraci) is, predominantly, a bone with a superficial ornament of dentine tubercles surrounded by pores leading to flask-shaped crypts (ampullae). However, despite the extensive bone present in these early dermal skeletons, damage was repaired almost exclusively with dentine. Consolidation of bone, by dentine invading and filling the vascular spaces, was previously recognized in Psammolepis and other heterostracans but was associated with ageing and dermal shield wear (reparative). Here, we describe wound repair by deposition of dentine directly onto a bony scaffold of fragmented bone. An extensive wound response occurred from massive deposition of dentine (reactionary), traced from tubercle pulp cavities and surrounding ampullae. These structures may provide the cells to make reparative and reactionary dentine, as in mammalian teeth today in response to stimuli (functional wear or damage). We suggest in Psammolepis, repair involved mobilization of these cells in response to a local stimulatory mechanism, for example, predator damage. By comparison, almost no new bone is detected in repair of the Psammolepis shield. Dentine infilling bone vascular tissue spaces of both abraded dentine and wounded bone suggests that recruitment of this process has been evolutionarily conserved over 380 Myr and precedes osteogenic skeletal repair.

Structural design and mechanical behavior of alligator (Alligator mississippiensis) osteoderms

Alligator is a well-adapted living fossil covered with dorsal armor. This dermal shield consists of bony plates, called osteoderms, interconnected by sutures and non-mineralized collagen fibers, providing a dual function of protection and flexibility. Osteoderm features a sandwich structure, combining an inner porous core and an outer dense cortex, to offer enhancements for stiffness and energy absorbance. In this study, we investigated the multi-scale structure and mechanical behaviors of the American alligator (Alligator mississippiensis) osteoderm. Microcomputed tomography was applied to reveal the complex neurovascular network. Through the observation under optical and scanning electron microscopes, the osteoderm was found to consist of woven bone in the dorsal region and lamellar-zonal bone in the ventral region. Nanoindentation and compressive tests were performed to evaluate the mechanical properties of osteoderms. The varying mineral contents and porosity result in a graded mechanical property: a hard and stiff dorsal cortex gradually transform to a more compliant ventral base. Three protective mechanisms optimized for alligator osteoderms were proposed and elucidated.

Description of the adult skeleton and developmental osteology of the hyperossified horned frog, Ceratophrys cornuta (Anura:Leptodactylidae).

The adult skeleton and tadpole chondrocranium of the leptodactylid frog, Ceratophrys cornuta (Ceratophryinae), are described in detail, including the ontogenetic development of the chondrocranium and the ossification sequence of the skeleton. The chondrocranium of the carnivorous larvae is unique in lacking a frontoparietal fontanelle and possessing a complete dorsal roof of cartilage. Furthermore, the chondrocranium is extremely robust, particularly those elements involved in the feeding mechanism; these include large palatoquadrate cartilages, stout Meckel's, supra-, and infrarostral cartilages, and short, wide, cornua trabeculae. The chondrocranium of C. cornuta resembles that described for Ceratophrys cranwelli, but differs from the chondrocrania reported for the species of Lepidobatrachus. The large adult skull is hyperossified; most elements are fused into a single unit, and nearly all dermal elements are ornamented, casqued, and co-ossified. Calcification is present in nearly every cartilaginous element of the skeleton in larger (older) adults. Several osteological characters previously used in ceratophryine systematics, such as the otic ramus of the squamosal and the columella, are reassessed. Contrary to previous reports, the ossified, dorsal dermal shield above the vertebral column in many ceratophryine anurans is absent in C. cornuta. With few exceptions, the ossification sequence relative to metamorphosis is consistent with those that are known for other anurans. The squamosal arises from three distinct centers of ossification, including an otic element. The frontoparietal arises from two centers of ossification that fuse early in development. A robust postorbital arch is formed primarily by the otic flange of the frontoparietal, which articulates laterally with the medial border of the otic ramus of the squamosal. Changes in the timing of development, or heterochrony, are involved with the evolution of the unusual skull and skeleton of ceratophryine frogs.

The development of a dermal shield in impala

The presence of a dermal shield in adult male impala is related to their righting style. The shield protects the area where blows are most likely to fall. Its development is compared with their social and behavioural development. It is suggested that the shield has, or has had, adaptive value.

The Evolution of Horn-Like Organs

Hornlike organs evolved independently in a number of mammalian families. Though these organs assumed great diversity they did evolve into several general functional types. A short review of the structure and development of hornlike organs is given. Some views on horn function and evolution are critically discussed. The evolution of hornlike organs is visualised as follows: In primitive large mammals the head blow became effective as a fighting form due to increased mass and inertia of the heads. Some forms grasped this potential. Combats were carried out from the broadside while opponents delivered head blows on each others body. Skull protuberances now became adaptive. Concurrently, defensive mechanisms evolve, decreasing the effectiveness of these protuberances. Foremost among them is a thick, heavy hide or specialised dermal shield. These adaptive syndromes gave no impetus towards larger and more complex horns. This impetus arose with the appearance of a new method of defense - catching the opponent's blows with the horned head. This leads to the evolution of heavy skulls and horns capable of catching and holding the opponent's head. The target area of attack remained the body. Frontal engagements resulted from the opponents' attempts to control each others horned head. It is shown that bovids and suids followed similar evolutionary roads in their mode of combat. The tusks of the suidae fulfill the same function and were subject to similar selection as the horns of short horned bovids. Thus Sus and Oreaisinos, and Bos and Phacochoerus are entirely similar in their mode of combat, hornlike organs and defense mechanisms. The primitive frontal engagement gave rise to two different modes of combat, ramming and wrestling. The wrestlers evolved complex horns, whose function is to bind opponents together, thus allowing them to develop full strength wrestling and pushing matches. The rammers evolved a heavy skull and heavy horns. A trend towards cephalisation of display organs can be detected in ruminants and suids. Display organs are interpreted as attention guiding adaptations. It is shown that the distribution of display organs correlates with the display. A broadside body display is found in ruminants with a diffuse distribution of display organs. If display organs are cephalised, the broadside display disappears. This is found in the North American wild sheep. Display centers about one heavy, very large horns of males. Next it is verified that the rams horns are display organs. It is shown, the horn display is the prerogative of the dominant; sheep can differentiate horn size classes; large horned males enjoy reproductive advantage; horn size carries a priori dominance status. Hence horns evolved to function as weapons inflicting damage; as defense organs shielding their owner; as binding organs allowing opponents a secure lock in battle; as display organs having an a priori intimidating effect on certain conspecifics. Some species of ruminants possess hornlike organs which change progressively in size with the age of their owner. It is postulated that, as in North American wild sheep, horn size parallels dominance order. Conspecifics learn to associate horn size and agonistic potential of opponents. Hence horn size signals to conspecifics predictable social relationships. In large cervids from temperate and cold climates this system must save stored energy during the rut, which males need for the severe winter. Graded horn sizes allow wild sheep to live in an open society in which strange individuals may meet and fit into the dominance hierarchy with a minimum of combat. Graded horn sizes hence serve as rank indicators. It is next made plausible that territories in ruminants function in conjunction with displaying males as rank indicators. Outside the territories males mix freely. Hence graded horn sizes and territoriality may enable open societies, by acting as communication mechanisms and creating a predictable, social environment. A hypothesis is presented explaining the selective forces for the evolution of display. It predicts that, antagonistic selection forces shape display and display-organs, that a potential exists for rapid spread in a population of differences in behaviour and morphology, that intimidation mechanisms evolved for all sensory pathways, that external appearance af animals changes more repidly than their primary adaptations.

Note on the discovery of the dermal shield in Megatheroid animals

"Note on the discovery of the dermal shield in Megatheroid animals." Annals and Magazine of Natural History, 18(104), p. 137