Wiley

The Anatomical Record

Published by Wiley and American Association for Anatomy

Online ISSN: 1932-8494

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Print ISSN: 1932-8486

Disciplines: Anatomy & physiology

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419 reads in the past 30 days

Time‐scaled phylogenetic relationships of the main lineages presented in this review. Relevant references for each group are cited throughout this work. Dotted lines represented relationships and phylogenetic placements that are still heavily debated. Thicker lines represent the known stratigraphic extent of the diverse clades. Shape outlines are taken from Phylopic (www.phylopic.org).
Hypothetical reconstitution of Terrestrisuchus gracilis. Taken from Irmis et al. (2013, fig. 5). Scale bar is 10 cm.
Reconstruction of Protosuchus richardsoni, based on AMNH 3024. Modified from Colbert and Mook (1951) (fig. 4). Scale bar is 5 cm.
Skeletal reconstruction of Araripesuchus gomesii (AMNH 24450). Modified from Hecht (1991). Scale bar is 5 cm.
Skeleton of Montealtosuchus arrudacamposi (MPMA‐16‐0007/04) from the Turonian–Santonian of Adamantina Formation (Brazil). (a) Dorsal view and (b) ventral view. Modified from Tavares et al. (2017) (fig. 1). Scale bar is 5 cm.

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A review of the non‐semiaquatic adaptations of extinct crocodylomorphs throughout their fossil record

November 2024

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422 Reads

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Aims and scope


The Anatomical Record: Advances in Integrative Anatomy and Evolutionary Biology publishes new discoveries in the morphological aspects of evolutionary, molecular, cellular, and systems biology. The journal focuses on major new findings in functional morphology, paleontology, neurology, histology, and the anatomical consequences of gene disruption, activation, or over expression. It also recognizes the importance of descriptive studies in contemporary research, particularly when framed in the context of experimental models or questions.

Recent articles


Variation and development of the turtle chondrocranium, with a description of the common musk turtle (Sternotherus odoratus, Kinosternidae, Cryptodira, Testudines)
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December 2026

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609 Reads

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2 Citations

Based on histological cross sections, the chondrocranium of the common musk turtle (Sternotherus odoratus) was reconstructed, described, and compared to other turtles. It differs from that other turtle chondrocrania by possessing elongated, slightly dorsally orientated nasal capsules with three dorsolateral foramina, which might be homologous to the foramen epiphaniale, and by having an enlarged crista parotica. Additionally, the posterior part of the palatoquadrate is more elongated and more slender than in other turtles, while its ascending process is connected to the otic capsule by appositional bone. The proportions of the chondrocranium were also compared to those of “mature” chondrocrania of other turtle species in a Principal Component Analysis (PCA). Other than expected, the S. odoratus chondrocranium is not similar in proportions to those of chelydrids, the closest related species in the sample. The results indicate to differences in the proportions among larger turtle clades (e.g., Durocryptodira, Pleurodira, and Trionychia). S. odoratus is an exception to this pattern since it shows elongated nasal capsules similar to the trionychid Pelodiscus sinensis. A second PCA comparing the chondrocranial proportions of multiple developmental “stages” mostly shows differences between trionychids and all other turtles. S. odoratus is again similar to trionychids along PC1, but its proportions are the most similar along PC2 and PC3 to older “stages” of americhelydians, including the chelydrid Chelydra serpentina, which is related to chondrocranium height and quadrate width. We discuss potential ecological correlations of our findings mirrored in late embryonic stages.


Cortico‐cortical connectivity of the somatosensory cortex of the agouti: Topographical organization and evolutionary implications

December 2024

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49 Reads

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Marco Aurelio M. Freire

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Understanding patterns of cortico‐cortical connections in both frequently and infrequently studied species advances our knowledge of cortical organization and evolution. The agouti (Dasyprocta aguti, a medium‐size South American rodent) offers a unique opportunity, because of its large lissencephalic brain and its natural behaviors, such as gnawing and hiding seeds, that require bimanual interaction while sitting on its hindlimbs and aligning its head to receive images of the horizon on the retinal visual streak. There have been no previous studies of the intrinsic and extrinsic ipsilateral projections of the agouti's primary somatosensory cortical area (S1). In the present study, we utilized biotinylated dextran (BDA) anatomical tract‐tracer injections combined with microelectrode electrophysiological mapping, correlated with analysis of cytochrome oxidase (CO) histochemical staining, to investigate the ipsilateral corticocortical connectivity of the agouti's S1. By injecting BDA into electrophysiologically identified regions within the S1, we revealed ipsilateral intrinsic connections, as well as connections with cortical areas rostral and caudal to S1, and homotopic labeling in the second somatosensory cortical area (S2). In addition, we identified a focal cluster of labeled axons and axonal terminals adjacent to the rhinal fissure, tentatively named the parietal rhinal area (PR). The analysis of CO reactivity allowed delineation of the boundaries and subdivisions of S1, as well as the locations and limits of primary auditory and visual areas. These findings provide support for the notion of a similar pattern of somatosensory cortical organization and connectivity across mammalian species.


The velar chord and dynamic integration of the gular valve in crocodylians

December 2024

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33 Reads

Crocodylians evolved a unique gular valve that is capable of creating a water‐tight seal between the oral and pharyngeal cavities, allowing the animal to safely submerge with an open mouth. The gular valve has traditionally been described as consisting of two separate parts: an active mobile ventral portion (consisting of the tongue and portions of the hyolingual apparatus) and a dorsal portion, which is a static fold on the hard palate (often termed the palatal velum). The results of the present study argue that the two portions of the gular valve are functionally integrated, not separate, and that the dorsal portion (herein the dorsal gular fold) is a dynamic element the shape and tension of which are influenced by active and passive forces. Using gross dissection, histology, and DiceCT, the present study documents a previously underscribed component of the gular valve, the velar chord, which links the hyolingual apparatus to the dorsal gular fold, functionally integrating the two halves of the gular valve. Through endoscopic videography and a variety of manipulations on living crocodylians, this study demonstrates that changes in the tension on the velar chord directly alter the shape and tension of the dorsal gular fold. The shape changes observed in the dorsal gular fold could be accommodated by a shallow depression in the ventral surface of the palatine bones, herein termed the velar fossa. The velar fossa is a prominent feature of Alligator mississippiensis and was observed in other crocodilians; however, a survey of living and fossil crocodylians demonstrated that the velar fossa is not a universal feature in this clade. Understanding the functional linkage between the dorsal and ventral portions of the gular valve has implications beyond the dive reflex of crocodylians, since active manipulation of the dorsal gular fold likely plays a role in a variety of behavioral and physiological processes such as deglutition and vocalization.


Cranial morphology and phylogenetic reassessment of Barreirosuchus franciscoi (Crocodylomorpha, Notosuchia), a Peirosauria from the Late Cretaceous of Brazil

December 2024

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103 Reads

With nearly 30 living species of relatively similar ecological traits, Crocodylomorpha is represented today by only a small fraction of its past diversity. The well‐documented crocodylomorph fossil record has revealed more than 500 taxa, with much higher ecological and morphological diversity than their extant counterparts. An example of such astonishing diversity is the Late Cretaceous rocks of the Bauru Group (southeast Brazil), from which numerous taxa are known, belonging to the clade Notosuchia. These were predominantly terrestrial taxa, some of which exhibited traits associated with omnivorous or even herbivorous feeding behaviors, such as Sphagesauridae, whereas others were adapted to a carnivore diet, such as Baurusuchidae and Peirosauridae. Among these is Barreirosuchus franciscoi, originally described as a neosuchian (Trematochampsidae) but later interpreted as a peirosaurid notosuchian. Even though included in recent morphological and phylogenetic analyses, B. franciscoi still lacked a more detailed description. Here, we provide an in‐depth description of the cranial elements of B. franciscoi, using data from computed tomography and a broad sample of comparative material, including living and fossil crocodylomorphs. Also, the neuro‐cavities, including the endocast, nasopharyngeal duct, and the olfactory region, were digitally reconstructed. Finally, a new phylogenetic analysis recovered B. franciscoi nested within Peirosauria, forming the Itasuchidae clade with other potentially semiaquatic species: Rukwasuchus yajabalajekundu, Pepesuchus deiseae, and Itasuchus jesuinoi. The morphological and phylogenetic reassessment of B. franciscoi indicates a semiaquatic form, highlighting the ecological diversity of notosuchians from the Bauru Group as well as the capacity of notosuchians to explore a myriad of environments.


Tracking points in x‐ray fluoroscopy image during respiration. A coordinate system (X–Y) was used as a reference for distance calculations. After the surfaces of each structure were labeled (black lines), the movements of the pharyngeal wall (PW, green dot), the tongue base (TB, pink dot), soft palate (SP, red dot), and the epiglottis (EP, blue dot) were each traced individually during inspiration, expiration, and pause between respiratory cycles. The cross at the lower left side indicates orientations of movements in the coordinate system. One of the eight 2 mm sonocrystals (black dots with wire shadows), previously implanted in the tongue base, was randomly used for calibration. NS, nasal cavity; VP, velopharynx. Scale: 10 mm.
Tracking points in x‐ray fluoroscopy image during chewing and swallowing. Distances and directions between SP, TB, EP, and PW (structural and inter‐structural) during both chewing cycles and swallowing episodes were measured. The bolus was also labeled to trace its movement from anterior to posterior and during swallowing episodes. Refer to Figure 1 for all captions and orientations.
Respiratory movements in the coordinate system for inspiration (a), expiration (b), pause (c). EP, epiglottis; PW, pharyngeal wall; SP, soft palate; TB, tongue base. Averages 10–12 respiratory cycles (20 s) per pig were included in the construction of these graphs. The arrows indicate changes in direction for each structure. Units in millimeters.
Early chewing movements in the coordinate system for pre‐feeding (a), jaw opening (b), jaw closing/power stroke (c), soft palate (orange, SP), epiglottis (gray, EP), tongue base (yellow, TB) and pharyngeal wall (blue, PWx, PWy). Averages 12–13 chewing cycles per pig were included in the construction of these graphs. Pre‐feeding movements were recorded while the minipig was awake without eating. The arrows indicate changes in direction for each structure. Units in millimeters.
Late chewing/pre‐swallowing movements and swallowing in the coordinate system for jaw opening (a), jaw closing/power stroke (b), and swallowing (c). Refer to Figure 4 for all captions and descriptions. Units in millimeters.
Spatial relationships of oropharyngeal structures during respiration, chewing, and swallowing

December 2024

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5 Reads

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1 Citation

Spatial relationships between oropharyngeal structures and their coordinated dynamics ensure proper adaptations in functions such as respiration, chewing, and swallowing. Thus, the aim of this study is to analyze spatial changes in the normal oropharynx during respiration and mastication. For this purpose, eight 7–8 months old Yucatan minipigs, four of each sex were used. X‐ray fluoroscopy was recorded with the field of view focused on the oropharyngeal region. The x‐ray video clips showing respiratory cycles and masticatory sequences were digitized and traced. Points to be digitized and traced were located on the soft palate, epiglottis, tongue base, and pharyngeal wall. An X–Y coordinate system was established to trace distances and directions of each structure (structural), and between structures (inter‐structural) during phases of respiration, chewing stages, and swallowing episodes. During respiration, the soft palate showed the largest X–Y movements with the largest distance change (1.32 ± 0.64 mm). In contrast, distance changes in the tongue base were significantly smaller (0.32 ± 0.21 mm, p < 0.05). Notably, during chewing the tongue base and epiglottis showed major changes in distance and direction. Similarly, during swallowing the tongue base showed the largest changes (2.94 ± 1.28 mm) followed by the pharyngeal wall and epiglottis. Thus, although coordinated, each pharyngeal structure plays specific roles. Understanding of these spatial and specific dynamics in different oropharyngeal structures would provide the baseline to analyze the potential mechanisms of various swallowing and breathing disorders such as dysphagia and obstructive sleep apnea.


Quantitative assessment of masticatory muscles based on skull muscle attachment areas in Carnivora

November 2024

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62 Reads

Masticatory muscles are composed of the temporalis, masseter, and pterygoid muscles in mammals. Each muscle has a different origin on the skull and insertion on the mandible; thus, all masticatory muscles contract in different directions. Collecting in vivo data and directly measuring the masticatory muscles anatomically in various Carnivora species is practically problematic. This is because some carnivorans can be ferocious, rare, or even extinct. Consequently, the most practical method to collect data on the force generated by the masticatory muscle is to estimate the force based on skulls. The physiological cross‐sectional area (PCSA) of each masticatory muscle, which correlates to the maximum force that can be produced by a muscle, was quantified. Using computed tomography, we defined the three‐dimensional measurement area for 32 carnivoran species based on the origin and insertion of masticatory muscles specified by observable crests, ridges, and scars. Subsequent allometric analysis relating the measurement area on skull surface to the PCSA for each masticatory muscle measured in fresh specimens revealed a strong correlation between the two variables. This finding indicates that within Carnivora, an estimation of absolute masticatory muscle PCSA can be derived from measurements area on skull surface. This method allows for the use of cranial specimens, housed in museums and research institutions, that lack preserved masticatory muscles in quantitative studies involving masticatory muscle PCSA. This approach facilitates comprehensive discussions on the masticatory muscle morphology of Carnivora, including rare and extinct species.


A review of the non‐semiaquatic adaptations of extinct crocodylomorphs throughout their fossil record

November 2024

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422 Reads

Crocodylomorphs constitute a clade of archosaurs that have thrived since the Mesozoic until today and have survived numerous major biological crises. Contrary to historic belief, their semiaquatic extant representatives (crocodylians) are not living fossils, and, during their evolutionary history, crocodylomorphs have evolved to live in a variety of environments. This review aims to summarize the non‐semiaquatic adaptations (i.e., either terrestrial or fully aquatic) of different groups from different periods, highlighting how exactly those different lifestyles are inferred for those animals, with regard to their geographic and temporal distribution and phylogenetic relationships. The ancestral condition for Crocodylomorpha seems to have been a terrestrial lifestyle, linked with several morphological adaptations such as an altirostral skull, long limbs allowing a fully erect posture and a specialized dentition for diets based on land. However, some members of this clade, such as thalattosuchians and dyrosaurids display adaptations for an opposite, aquatic lifestyle, interestingly inferred from the same type of morphological observations. Finally, new techniques for inferring the paleobiology of those extinct animals have been put forward in the last decade, appearing as a complementary approach to traditional morphological descriptions and comparisons. Such is the case of paleoneuroanatomical (CT scan data), histological, and geochemical studies.


Functional and phylogenetic signals in the pectoral girdle of Thalattosuchia and Dyrosauridae (Crocodylomorpha)

November 2024

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162 Reads

Crocodylomorphs have colonized various environments from fully terrestrial to fully aquatic, making it an important clade among archosaurs. A remarkable example of the rich past diversity of Crocodylomorpha Hay, 1930 is the marine colonization undergone by several crocodylomorph lineages, particularly Thalattosuchia Fraas, 1901 during the Early Jurassic–Early Cretaceous, and Dyrosauridae de Stefano, 1903 during the Late Cretaceous–Early Eocene. Thalattosuchia represents the most impressive and singular marine radiation among Crocodylomorpha, occupying various ecological niches, before enigmatically disappearing in the Cretaceous. Dyrosauridae, on the other hand, is known for surviving the end‐Cretaceous mass extinction in abundance but subsequently vanished. The evolutionary path undertaken by crocodylomorphs into the aquatic environments and the reasons for their disappearance outside marine extinction events during the Mesozoic remains a mystery. Despite a well‐preserved fossil record, attention has primarily centered on craniodental adaptations, overlooking the swimming‐related adaptations recorded in the postcranial skeleton. This research primarily involves a comprehensive examination of the pectoral girdle of the most representative members of Thalattosuchia and Dyrosauridae, highlighting their evolutionary trajectories over time. Additionally, this work aims to test the phylogenetic signal residing in the postcranial anatomy of Crocodylomorpha. As such, the most recent and complete Crocodylomorpha phylogenetic dataset has been repurposed: 42 new postcranial characters have been added and several others have been revised to address our phylogenetic question. We stress that postcranial anatomy constitutes an important tool supply to better understand the relations of extinct crocodyliforms, but also offers insights on their development, ecology, and biomechanics.


Synapsids and sensitivity: Broad survey of tetrapod trigeminal canal morphology supports an evolutionary trend of increasing facial tactile specialization in the mammal lineage

November 2024

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143 Reads

The trigeminus nerve (cranial nerve V) is a large and significant conduit of sensory information from the face to the brain, with its three branches extending over the head to innervate a wide variety of integumentary sensory receptors, primarily tactile. The paths of the maxillary (V2) and mandibular (V3) divisions of the trigeminus frequently transit through dedicated canals within the bones of the upper and lower jaws, thus allowing this neuroanatomy to be captured in the fossil record and be available to interpretations of sensory ability in extinct taxa. Here, we use microCT and synchrotron scans from 38 extant and fossil species spanning a wide phylogenetic sample across tetrapods to investigate whether maxillary and mandibular canal morphology can be informative of sensory biology in the synapsid lineage. We found that in comparison to an amphibian and sauropsid outgroup, synapsids demonstrate a distinctive evolutionary pattern of change from canals that are highly ramified near the rostral tip of the jaws to canals with increasingly simplified morphology. This pattern is especially evident in the maxillary canal, which came to feature a shortened infraorbital canal terminating in a single large infraorbital foramen that serves as the outlet for branches of V2 that then enter the soft tissues of the face. A comparison with modern analogues supports the hypothesis that this morphological change correlates to an evolutionary history of synapsid‐specific innovations in facial touch. We interpret the highly ramified transitional form found in early nonmammalian synapsids as indicative of enhanced tactile sensitivity of the rostrum via direct or proximal contact, similar to tactile specialists such as probing shorebirds and alligators that possess similar proliferative ramifications of the maxillary and mandibular canals. The transition toward a simplified derived form that emerged among Mid‐Triassic prozostrodont cynodonts and is retained among modern mammals is a unique configuration correlated with an equally unique and novel tactile sensory apparatus: mobile mystacial whiskers. Our survey of maxillary and mandibular canals across a phylogenetic and ecological variety of tetrapods highlights the morphological diversity of these structures, but also the need to establish robust form‐function relationships for future interpretations of osteological correlates for sensory biology.


The first ontogenetic model for non‐crocodylomorph loricatans, based on osteohistology of the ontogenetic series of Prestosuchus chiniquensis from the Middle Triassic of Brazil

November 2024

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107 Reads

Prestosuchus chiniquensis is an iconic non‐crocodylomorph loricatan from the Brazilian Triassic beds and the best‐known taxon, represented by several specimens. The completeness and preservation of its skeleton make it a valuable taxon for paleobiological studies. We explore the microstructure of bone tissues of appendicular elements and ribs of three specimens of Prestosuchus to access a variety of aspects of its paleobiology, such as histovariability, ontogeny, and growth patterns. Integrating our data and other osteohistologically studied P. chiniquensis specimens, we proposed for the first time an ontogenetic model for non‐crocodylomorph loricatans. The model encompasses six distinct age classes (I–VI) that allow us to infer the growth patterns of P. chiniquensis and possibly expand to other phylogenetically close taxa. During early ontogeny (age classes I–II), sustained fast growth was maintained by a fibrolamellar complex. In mid ontogeny (age classes III–IV), highly vascularized parallel‐fibered bone predominates, suggesting intermediary growth rates. A change for a poorly vascularized parallel‐fibered/lamellar bone would mark the attainment of sexual (age classes IV–V) and skeletal maturity, comprising the age class VI. An external fundamental system (EFS) present in the outermost cortex is the main histological feature that characterize the age class VI. Major histovariability features are present between appendicular bones and ribs of skeletally immature individuals. The most prominent of them is the presence of fibrolamellar complex and highly vascularized parallel‐fibered bone in appendicular bones and poorly vascularized parallel‐fibered bone in ribs. In advanced ontogenetic stages, the histovariability between appendicular bones and ribs tends to be minor. Our data also support previous hypothesis of the presence of one new taxon among the specimens assigned to P. chiniquensis, increasing the diversity of non‐crocodylomorph loricatans. The new taxon, represented by the specimen UFRGS‐PV‐0152‐T, awaits a formal anatomical description. Our study advances the preliminary understand of the ontogeny and growth patterns of non‐crocodylomorphs loricatans and Pseudosuchia as a whole.


Sniffing out morphological convergence in the turbinal complex of myrmecophagous placentals

November 2024

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121 Reads

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1 Citation

The length of the snout in mammals has important evolutionary consequences for the functional systems housed within the rostrum. However, whether increased snout lengths lead to expanded olfactory performance has rarely been examined. Here, we investigate inner rostral function among 10 species of myrmecophagous (ant‐ and/or termite‐eating) placental mammals and 10 closely related species. We use nondestructive computed tomography scanning methods to characterize inner rostral function based on the underlying morphology of the turbinal bones in the nasal cavity. Three approaches were chosen to address this question, including the quantification of functional turbinal surface area, the quantification of functional turbinal three‐dimensional complexity, and geometric morphometrics. By including non‐model species from several different mammalian orders, we were able to extend the discussion surrounding turbinal homologies to comparisons across mammals. Our results show no increased olfactory function in all myrmecophagous species relative to their sister taxa, which suggests that there is no trade‐off for increased olfactory capabilities in myrmecophagous species with elongated snouts. We found no evidence of convergence in turbinal morphology among all five myrmecophagous lineages. However, we found evidence of morphological convergence in the turbinals between the giant armadillo and the aardvark, suggesting a more complex interplay between the evolution of turbinal morphology and ecological correlates. While myrmecophagy alone may not be a strong enough ecological signal to overcome phylogenetic and developmental constraints, we suggest that this might be the case at the intersection of this dietary specialization with a primarily underground lifestyle where odorants may be difficult to detect.


Anatomy and ontogeny of the “carnivorous aetosaur”: New information on Coahomasuchus kahleorum (Archosauria: Pseudosuchia) from the Upper Triassic Dockum Group of Texas

November 2024

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160 Reads

A newly referred specimen of Coahomasuchus kahleorum (TMM 31100‐437) from the lower part of the Upper Triassic Dockum Group of Texas preserves much of the skeleton including the majority of the skull. Introduced in the literature in the 1980s as the “carnivorous aetosaur”, TMM 31100‐437 bears recurved teeth that previously were considered unique among aetosaurs. The small size of the individual led to speculation that it represents a skeletally immature individual that retains a plesiomorphic dentition for Archosauromorpha. We provide a detailed evaluation of the anatomy and phylogenetic relationships of this specimen. Apomorphies of the osteoderms and braincase support the referral of the specimen to C. kahleorum. Histological analysis of the femur demonstrates that TMM 31100‐437 does not represent a juvenile form of another known aetosaur. Thus, TMM 31000‐437 provides another case demonstrating that aetosaur species spanned a wide range of maximum body sizes, from approximately 1.5 m to over 5 m in length. Reanalysis of the type specimen of C. kahleorum, along with information from TMM 31000‐437, demonstrates that the lateral osteoderms are not autapomorphic as previously described and have distinct lateral and medial flanges as well as a dorsal eminence. Overall, this specimen provides key details regarding body size and diet in an early occurring aetosaur.


Morphological characteristics of the unique periodontal structure in dolphins

November 2024

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60 Reads

Although toothed whales have dentition peculiar to mammals, little attention has been paid to the periodontal tissues that support these characteristic teeth. In this study, we clarified the anatomical characteristics of the periodontal tissue in several species of Delphinidae through three‐dimensional observation using micro‐computed tomography, histological observations using decalcified sections, and immunohistochemical analysis. The results indicated that the teeth and the periodontal tissues of dolphins are morphologically unique among mammals. The alveolar bone was both crude and spongy. The lamina dura, a radiopaque line observed in the alveolar bone of common mammals, was thin in dolphins, and the teeth were attached to the trabeculae with the periodontal ligament (PDL). The alveolar sockets were massive for the size of the teeth. The PDL, a collagen fiber that fills the periodontal space, was well‐developed and peculiarly divided into two layers. The inner layer fibers radially spread out from the cementum, similar to the PDL in common mammals. However, the outer layer fibers penetrate the spongy bone in a complicated manner. The interstitial space between the inner and outer layers contained nerve fiber bundles that were thicker than those found in the PDL of other mammals. Sensory receptor‐like structures were observed at the terminal ends of the nerve fibers. These findings indicated that the dolphin PDL is more sensitive to dental stimuli than those of other mammals, suggesting that the dolphin dentition plays a functional role as a sensory receptor, similar to tactile hair.


Mandible composition and properties in two selected praying mantises (Insecta, Mantodea)

November 2024

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153 Reads

Insects process their food with their cuticle‐based mouthparts. These feeding structures reflect their diversity and can, in some cases, showcase adaptations in material composition, mechanical properties, and shape to suit their specific dietary preferences. To pave the way to deeply understand the interaction between mouthparts and food and to determine potential adaptations of the structures to the food, this study focuses on the mandibles of two praying mantis species. Gongylus gongylodes feeds mainly on Diptera, and Sphodromantis lineola forages on larger prey. Employing scanning electron microscopy, the mandibular morphologies were analyzed. The degree of the cuticle tanning was tested using confocal laser scanning microscopy. Furthermore, the contents of transition and alkaline earth metals in the mandible cuticle were studied using energy‐dispersive X‐ray spectroscopy and the mechanical properties tested by nanoindentation. We found that S. lineola mandibles show pronounced gradients of Young's modulus and hardness from the basis to the tip, which might be an adaptation against high stresses during biting and chewing. G. gongylodes, in contrast, did not show pronounced gradients, which may indicate that there is less stress involved in feeding—necessary to test in future studies. The mechanical properties of manidibles in both species are related to the degree of cuticle tanning but also positively correlate with the content of magnesium. These findings enrich our understanding of insect cuticle biology but also present new sets of data on praying mantis structures.


Review of respiratory anatomy adaptations in whales

November 2024

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246 Reads

Whales (cetaceans, including dolphins and porpoises) are superbly adapted to life in water, but retain vestiges of their terrestrial ancestry, particularly the need to breathe air. Their respiratory tract exhibits many differences from their closest relatives, the terrestrial artiodactyls (even toed ungulates). In this review, we describe the anatomy of cetacean respiratory adaptions. These include protective features (e.g., preventing water incursions during breathing or swallowing, mitigating effects of pressure changes during diving/ascent) and unique functions (e.g., underwater sound production, regulating gas exchange during the dive cycle).


Theoretical distances used to determine the bite force. The TMJ resembles the fulcrum, and the muscle moment arms (MA) are vertical to the direction of the muscle force. The bite point moment arm (MAbite) connects each bite point on the first, second, and third molar and fulcrum.
Principal component analysis (PCA), separately for each size class. (a) Size class 1, (b) Size class 2, (c) Size class 3, (d) vectors of included parameters.
Boxplots of exemplary DMTA parameters from the functional categories complexity, height, and volume. (a) Sdr (developed interfacial ratio), (b) metf (mean depth of furrows), (c) Sz (maximum height), (d) Vm (material volume). *p = 0.05; **p = 0.01.
Correlation between bite force of all tooth positions and exemplary DMTA parameters per size class. (a) Sdr (developed interfacial ratio) shows no significant correlation; (b) metf (mean depth of furrows) shows a significant correlation in size class 1; (c) Sz (maximum height) shows a significant correlation in size class 1; (d) Vm (material volume) shows significant correlations in size classes 1 and 2.
Optimized, simplified patterns found for DMTA gradients along the upper tooth row for the three size classes. (a) Trend observed for several height and volume parameters. In size class 1, M1 and M2 show larger parameter values than M3, while for size class 2 and 3, tooth positions show similar parameter values. (b) Trend observed for height, complexity and slope parameters. In size class 1, M2 shows larger parameter values than M1 and M3. Size class 2 displays similar parameter values for all tooth positions, while a trend for increasing value scan be seen in size class 3.
Tooth eruption status and bite force determine dental microwear texture gradients in albino rats (Rattus norvegicus forma domestica)

October 2024

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84 Reads

Dental microwear texture analysis (DMTA) is widely applied for inferring diet in vertebrates. Besides diet and ingesta properties, factors like wear stage and bite force may affect microwear formation, potentially leading to tooth position‐specific microwear patterns. We investigated DMTA consistency along the upper cheek tooth row in young adult female rats at different growth stages, but with erupted adult dentitions. Bite forces for each molar (M) position were determined using muscle cross‐sectional areas and lever arm mechanics. Rats were categorized into three size classes based on increasing skull length. Maximum bite force increased with size, while across all size classes, M3 bite force was almost 1.4 times higher than M1 bite force. In size class 1, M1 and M2 showed higher values than M3 for DMTA complexity, height, and volume parameters, while in size class 3, M1 had the lowest values. Comparing the same tooth position between size classes revealed opposing trends: M1 and M2 showed, for most parameters, decreasing roughness and complexity from size class 1–3, while M3 displayed the opposite trend, with size class 1 showing lowest, and either size class 2 or 3 the highest roughness and complexity values. This suggests that as rats age and M3 fully occludes, it becomes more utilized during mastication. DMTA, being a short‐term diet proxy, is influenced by eruption and occlusion status changes. Our findings emphasize the importance of bite force and ontogenetic stage when interpreting microwear patterns and advise to select teeth in full occlusion for diet reconstruction.


Arrangements of intramembranous muscles of wings are influenced by body mass across the radiation of phyllostomid bats

October 2024

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114 Reads

Extensive research into bat flight mechanisms has highlighted the complex functional and evolutionary dynamics of their wing structures, yet the anatomical details of certain wing muscles remain elusive. In particular, the intramembranous plagiopatagiales proprii muscles, located within the plagiopatagium—an area of the wing lacking direct joint connections—exhibit remarkable variation across bat families. These muscles, which extend anteroposteriorly in macroscopic bundles, play a crucial role in wing stiffening, modulating membrane tension, and reducing wing curvature during flight. Since larger bats tend to have higher wing loading (WL; the ratio of body mass [BMa] to wing area) and may therefore experience increased patagial curvature and resultant drag, we hypothesized that body size significantly influences the evolutionary development of the plagiopatagiales proprii muscles. This study investigates the relationship between BMa and the morphology of the plagiopatagiales proprii in New World leaf‐nosed bats (Phyllostomidae), employing bivariate allometry, multivariate analysis, and comparative phylogenetic methods across 24 species from eight phyllostomid subfamilies. Our findings reveal a significant phylogenetic signal in muscle architecture, along with positive evolutionary allometry in muscle area. This suggests an adaptive increase in muscle size in larger species, likely to counterbalance the increased WL, reduce wing curvature, and minimize drag. This research enhances our understanding of the functional and adaptive morphological evolution of intramembranous wing muscles in phyllostomid bats, underscoring their evolutionary significance.


Nuclear parcellation of pontine catecholaminergic and cholinergic neurons in gray parrots and pied crow brains

October 2024

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24 Reads

Employing immunohistochemical procedures with antibodies raised against tyrosine hydroxylase (TH) and choline acetyltransferase we identified and mapped the locus coeruleus complex (LoC) and the pontine laterodorsal tegmental (LDT) and pedunculopontine tegmental (PPN) cholinergic nuclei in the brains of a Congo gray parrot, a timneh gray parrot, and a pied crow. The LoC and LDT/PPN are centrally involved in the regulation and generation of different sleep states, and as all birds studied to date show both REM and non‐REM sleep states, like mammals, we investigated whether these noradrenergic and cholinergic nuclei in the avian pons shared anatomical features with those in the mammalian pons. The LoC was parcellated into 3 distinct nuclei, including the locus coeruleus (A6), subcoeruleus (A7), and the fifth arcuate nucleus (A5), while distinct LDT and PPN nuclei were revealed. Several similarities that allow the assumption of homology of these nuclei between birds and mammals were revealed, including their location relative to each other and other structures within the pontine region, as well as a specific degree of topographical overlap of the noradrenergic and cholinergic neurons. Despite this, some differences were noted that may be of interest in understanding the differences in sleep between birds and mammals. Further anatomical and physiological studies are needed to determine whether these pontine nuclei in birds play the same role as in mammals, as while the homology is apparent, the functional analogy needs to be revealed.


Effects and phenotypic consequences of transient thyrotoxicosis and hypothyroidism at different stages of zebrafish Danio rerio (Teleostei; Cyprinidae) skeleton development

The effects and consequences of changes in thyroid hormones (THs) level are among the actively studied topics in teleost developmental and evolutionary biology. In most of the experimental models used, the altered hormonal status (either hypo‐ or hyperthyroidism) is a stable characteristic of the developing organism, and the observed phenotypic outcomes are the cumulative consequences of multiple TH‐induced developmental changes. Meanwhile, the influence of the transient fluctuations of TH content on skeleton development has been much less studied. Here, we present experimental data on the developmental effects and phenotypic consequences of transient, pharmacologically induced thyrotoxicosis and hypothyroidism at different stages of ossified skeleton patterning in zebrafish. According to the results, the skeleton structures differed in TH sensitivity. Some showed a notable shift in the developmental timing and rate, while other demonstrated little or no response to changes in TH content. The developmental stages also differed in TH sensitivity. We identified a relatively short developmental period, during which changes in TH level significantly increased the developmental instability and plasticity, leading to phenotypic consequences comparable to those in fish with a persistent hypo‐ or hyperthyroidism. These findings allow this period to be considered as a critical developmental window.


Bone microstructure analyses in ontogenetic series of Mesosaurus tenuidens from the early Permian of Brazil

Osteohistological evidence is widely used to infer paleobiological traits of fossil vertebrates, such as ontogeny and growth rates. Mesosaurs, an enigmatic group of aquatic reptiles from the early Permian, are the most well‐known Paleozoic amniotes from Africa and South America. Their fossils are abundant in South America, ranging from the central‐west region of Brazil to the southernmost areas, as well as parts of Paraguay and Uruguay. In this contribution, we examined the bone microstructure of Mesosaurus tenuidens by analyzing thin sections of axial and appendicular elements of several specimens collected from various Brazilian sites. The microstructure of the bones showed minimal histological variability among elements, predominantly composed of parallel‐fibered tissues, indicating slow growth rhythm, along with increased bone density attributed to pachyosteosclerosis. The cortical area consists of poorly vascularized parallel‐fibered bone tissue, which was interrupted by multiple cyclical growth marks, some of them being supernumerary, suggesting a strong influence of seasonality. Moreover, the organization of growth marks suggests distinct life history trajectories among individuals collected from different outcrops, reflecting environmental heterogeneity throughout the basin. Internally, the endosteal domain exhibits greater vascularization compared to the cortices and frequently contained calcified cartilage. In the ontogenetic series, there was a progressive filling of the medullary region from small to large individuals. The presence of the External Fundamental System (a proxy indicating somatic maturity) was observed in femora and ribs, suggesting that determinate growth was already occurring in Permian mesosaurs and may not be an exclusive specialization of crown amniotes.


Identification of acinar injured zones in the pancreas of control and hypothyroid (Hypo) rabbits. (a) Zone without damage, (b) Zone 1: infiltration of immune cells, localization of vacuoles in acinar cells, and anisonucleosis (asterisks). (c) Zone 2: loss of acinar cells and replacement with connective tissue (black arrows). (d) Zone 3: neogenesis of adipose tissue substituting acinar cells (white arrows). Scale = 20 μm; acinus (ac). Quantification of superoxide dismutase (SOD, (e), catalase (CAT, f), TBARS test (g), antioxidant status (TAS, h), and conjugated dienes (i) in the pancreas of control and hypothyroid rabbits. Control and Hypo n = 6, except for control of MDA n = 4. Data are expressed as medians ± SD.
Quantification of immune cells (a and b), macrophages CD163+ (c–h), and immune cells FXRα+ in the pancreas (i–n) of control and hypothyroid (Hypo) female rabbits. Data are expressed as mean ± SEM. **p < 0.01, and ***p < 0.001. Large (L), medium (M), and small (S) islets. Scale = 50 μm. Acinus (ac) and islets (is). White arrows indicate immune cells positive to markers CD163 or FXRα. Asterisks indicate islet cells positive to markers.
Serum and pancreatic concentration of triacylglycerol (TAG; a and c) and total cholesterol (TC; b and d), and expression of LDLR in the pancreas (e) for control and hypothyroid (Hypo) female rabbits. Data are expressed as means ± SEM (n = 6; except pancreatic cholesterol in the control group n = 5). *p < 0.05. The LDLR expression is relative to ~90% of each Ponceaus Red stained lane of total proteins.
Expression of CYP51A1 (a), FXRβ (b), 3β‐HSD (c), and PPARβ/δ (d) in the pancreas of control and hypothyroid (Hypo) female rabbits. Data are expressed as means ± SEM (n = 6). *p < 0.05; ***p < 0.001. The CYP51A1, FXRβ, and 3βHSD expressions are relative to each Ponceaus Red stained lane. CYP51A1and FXRβ expressions were detected chemiluminescent signal using an analyzer, but for 3β‐HSD the signals were captured using photographic plates. Scale = 50 μm. Acinus (ac) and islets (is). Black arrow: Immune cell.
Pancreatic inflammation induced by hypothyroidism in female rabbits is associated with cholesterol accumulation and a reduced expression of CYP51A1, FXRβ, and PPARβ/δ

In women and animal models, hypothyroidism induces hypercholesterolemia, pancreatitis, and insulitis. We investigated whether lipids are involved in the effects of hypothyroidism in the pancreas. Control (n = 6) and hypothyroid (n = 6) adult female rabbits were used. We quantified serum and pancreatic triacylglycerol and total cholesterol levels, the oxidative and antioxidant status, and the expression of low‐density lipoprotein cholesterol receptor (LDLR) in the pancreas. Inflammation of the pancreas was evaluated by infiltration of immune cells positive to CD163 and α‐farnesoid receptor (FXRα). Other lipid players involved in both inflammation and insulin secretion of the pancreas, such as lanosterol 14‐α‐demethylase (CYP51A1), β‐farnesoid receptor (FXRβ), 3β‐hydroxysteroid dehydrogenase (3β‐HSD), and peroxisome proliferator‐activated receptor β (PPARβ/δ), were quantified. Groups were compared by t‐Student or U‐Mann–Whitney tests (p ≤ 0.05). Hypothyroidism induced hypercholesterolemia and a high cholesterol accumulation in the pancreas of female rabbits, without affecting oxidative or antioxidative status nor the expression of LDLR. The pancreas of hypothyroid females showed inflammation identified by a great infiltration of immune cells, macrophages CD163+, and loss of expression of FXRα+ in immune cells. Moreover, a reduced expression of CYP51A1, FXRβ, and PPARβ/δ, but not 3β‐HSD, in the hypothyroid pancreas was found. Pancreatitis and insulitis promoted by hypothyroidism may be related to the accumulation of cholesterol, lanosterol actions, and the activation of PPARβ/δ. All inflammatory markers evaluated in this study are related to glucose regulation, suggesting the link between hypothyroidism and diabetes.


Exploring elbow kinematics in the central bearded dragon (Pogona vitticeps) using XROMM: Implications for the role of forearm long‐axis rotation in non‐avian reptile posture and mobility

The functional morphology and kinematics of the elbow joint remain relatively understudied in squamates. Previous investigations of lizard elbow morphology and kinematics suggest long‐axis rotation (LAR) of the radius and ulna during stance allows the manus to remain pronated during forelimb retraction. Using XROMM (X‐ray Reconstruction of Moving Morphology), we explored the range of 3D movements and kinematics of the humerus, radius, and ulna in three adult male Central bearded dragons (Pogona vitticeps) during trackway walking. Our data indicate that the elbow joint of P. vitticeps experiences significant rotations in all three dimensions and that the radius and ulna adduct and rotate laterally on their long axes relative to the elbow joint and to one another during stance. These movements allow the distal ends of the radius and ulna to remain in a configuration necessary for manus pronation. These data support previous inferences that the radius and ulna of lizards move independently at the wrist joint. We suggest that independent LAR of the radius and ulna relative to the elbow joint and to one another may be an ancestral mechanism in lizards and perhaps more broadly across non‐avian reptiles.


First record of rhynchosaurs (Archosauromorpha: Rhynchosauria: Hyperodapedontinae) from the early Late Triassic Santacruzodon Assemblage Zone of the Santa Maria Supersequence, Brazil

October 2024

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125 Reads

Rhynchosauria is a group of extinct, exclusively Triassic, terrestrial, and herbivorous archosauromorphs, characterized by a peculiar maxillary‐mandibular apparatus. They reached global distribution during the Carnian, with the Hyperodapedontinae clade. The rhynchosaurian record from South America is included in the Ladinian‐?earliest Carnian Dinodontosaurus Assemblage Zone (Pinheiros‐Chiniquá Sequence, Brazil) and Tarjadia Assemblage Zone (Chañares Formation, Argentina), and for strictly Carnian Hyperodapedon Assemblage Zone (Lower Candelária Sequence, Brazil) and Ischigualasto Formation (Argentina). Here, we present the first record of Hyperodapedontinae from the Brazilian Early Carnian Santacruzodon Assemblage Zone (Santa Cruz Sequence). The most diagnostic elements belong to a nearly complete left pes, which presents a combination of hyperodapedontine traits. The presence of a second individual of cf. Hyperodapedontinae is based on a fragmentary, isolated left metatarsal IV. An isolated fragment of premaxilla could belong to the second specimen (based on preservation mode) or to a third individual. This new report fills a gap within the South American rhynchosaurian distribution, strengthening biostratigraphic correlation with other regions from Gondwana (i.e., Madagascar), where similar and coeval faunas are known. The inclusion of these specimens in a phylogenetic dataset resulted in low resolution results, due to missing data because postcranial characters for rhynchosaurs are still poorly explored.


Meningeal transcriptomics of adult male Downy Woodpeckers (Dryobates pubescens) and comparison species (Tufted Titmouse [Baeolophus bicolor]). (a) Schematic of experimental design. (b) Venn diagram displaying gene ortholog overlap and differentiation between woodpeckers and comparison species. (c) Principal components analysis (PCA) revealing distinct clusters for Woodpeckers and comparison species. (d) Similar expression levels of genes specific to each meningeal layer (padj > 0.05), suggesting comparable tissue quantities. MAV, meningeal and associated vasculature.
Differential gene expression in the meningeal and associated vasculature (MAV) tissue of adult male Downy Woodpeckers and comparison species (Tufted Titmouse [Baeolophus bicolor]). (a) Volcano plot illustrates genes with a fold change (i.e., Log2 fold change) of ≥1.5 and a false discovery rate (FDR) >0.1, highlighting 3,403 genes overrepresented in woodpeckers (red, labeled “Higher”) and 3,593 genes underrepresented (blue, labeled “Lower”) (b) Heatmap of the top 80 differentially expressed genes (DEGs), divided into the top 40 overexpressed and top 40 underexpressed genes based on log2 fold change across all samples. Genes are marked with asterisks to highlight those of interest for their roles in immune and stress response (red asterisk; “Immune”), glymphatic system, waste clearance, and cerebrospinal fluid (CSF) management (green asterisk; “Homeostatic”), and cytoskeletal, ciliary, and other structural functions (blue asterisk; “Structure”). (c–e) Tukey box‐and‐whisker plots with overlaid scatter plots highlighting significant DEGs identified in recent studies (2017–2024) as differentially expressed in the MAV or meninges due to head trauma, categorized by their roles in (c) immune response, (d) homeostatic processes, and (e) structural elements.
Pathway enrichment analysis focusing on the top Gene Ontology (GO) Molecular Functions (MF) and Biological Processes (BP) in differentially expressed genes. Red, green, and blue asterisk next to a term represent, “Immune,” “Homeostatic,” and “Structural” functions, respectively. (a) The top 15 GO terms by adjusted p‐value from the top 2,000 differentially expressed genes based on absolute value of log fold change (LFC). (b) The top 25 GO terms derived from the top 2,000 overrepresented genes based on LFC. (c) The top 25 GO terms derived from the top 2,000 underrepresented genes based on LFC.
Comparative analysis of meningeal transcriptomes in birds: Potential pathways of resilience to repeated impacts

The meninges and associated vasculature (MAV) play a crucial role in maintaining cerebral integrity and homeostasis. Recent advances in transcriptomic analysis have illuminated the significance of the MAV in understanding the complex physiological interactions at the interface between the skull and the brain after exposure to mechanical stress. To investigate how physiological responses may confer resilience against repetitive mechanical stress, we performed the first transcriptomic analysis of avian MAV tissues using the Downy Woodpecker (Dryobates pubescens) and Tufted Titmouse (Baeolophus bicolor) as the comparison species. Our findings reveal divergences in gene expression profiles related to immune response, cellular stress management, and protein translation machinery. The male woodpeckers exhibit a tailored immune modulation strategy that potentially dampens neuroinflammation while preserving protective immunity. Overrepresented genes involved in cellular stress responses suggest enhanced mechanisms for mitigating damage and promoting repair. Additionally, the enrichment of translation‐associated pathways hints at increased capacity for protein turnover and cellular remodeling vital for recovery. Our study not only fills a critical gap in avian neurobiology but also lays the groundwork for research in comparative neuroprotection.


Different views of extant elephant skulls. (a) Elephas maximus cranium with mandible, lateral view; (b) Loxodonta africana cranium, lateral view; (c) Loxodonta africana cranium, rostral view; (d) Loxodonta africana mandible, lateral view; (e) Elephas maximus cranium with mandible, rostral view; (f) Loxodonta africana cranium with mandible, rostral oblique view; and (g) hemisected Elephas maximus cranium with mandible, inner sagittal view showing narial opening and passageway into trunk and cranial air cells. Not to scale. cac, cranial air cells; cp, coronoid process; if, incisive fossae; iof, infraorbital foramen; mc, mandibular condyle; mr, mandibular ramus; ms, mandibular symphysis; na, narial aperture; np, nasal process; o, orbit; pt, premaxillary tusks; pts, premaxillary tusk sheath; tf, temporal fossa; za, zygomatic arch.
Musculature of the proboscis (trunk) and feeding apparatus in extant elephants. (a) Elephas maximus superficial (top) and deep (bottom) craniofacial musculature; (b) Loxodonta africana superficial craniofacial musculature (top) and muscles of the trunk in cross section (bottom); and (c) Loxodonta africana skull showing muscle attachment sites of m. temporalis (blue) and m. masseter (red). Not to scale. ln, m. lateralis nasi; m, m. masseter; ml, m. maxillo‐labialis; n, m. nasalis; nl, m. nasolabialis; pr, m. pars rimana; ps, m. pars supralabialis; rn, m. rectus nasi; rnd., dorsal fibers of rectus nasi; rnt, transverse fibers of m. rectus nasi; rnv, ventral fibers of m. rectus nasi; t, m. temporalis.
Skulls of earlier‐diverging proboscideans. (a) Moeritherium cranium with reconstructed mandible, lateral view; (b) Deinotherium cranium with mandible, lateral view (flipped image); (c) Deinotherium cranium with mandible, rostral view; (d) Palaeomastodon cranium with mandible, lateral view (flipped image); (e) Mammut cranium with mandible, lateral view; and (f) Mammut cranium with mandible, rostral oblique view. Not to scale. cp, coronoid process; if, incisive fossae; iof, infraorbital foramen; mc, mandibular condyle; mr, mandibular ramus; ms, mandibular symphysis; mt, mandibular tusks; na, narial aperture; np, nasal process; o, orbit; pt, premaxillary tusks; pts, premaxillary tusk sheath; sr, symphyseal ramus; tf, temporal fossa; za, zygomatic arch.
Skull of later‐diverging proboscideans. (a) Gomphotherium cranium with mandible, lateral view (flipped image); (b) Gomphotherium cranium with mandible, rostral view; (c) Archaeobelodon cranium with mandible, lateral view; (d) Platybelodon mandible, rostral oblique view; (e) Stegodon cranium, lateral view; and (f) Mammuthus cranium with mandible, rostral oblique view (flipped image). Not to scale. cp, coronoid process; if, incisive fossae; iof, infraorbital foramen; mc, mandibular condyle; mr, mandibular ramus; ms, mandibular symphysis; mt, mandibular tusks; na, narial aperture; np, nasal process; o, orbit; p, premaxilla; pt, premaxillary tusks; pts, premaxillary tusk sheath; sr, symphyseal ramus; ssg, suprasymphyseal groove; tf, temporal fossa; za, zygomatic arch.
Craniofacial musculature and feeding directionality in proboscideans. (a) Craniofacial muscle reconstructions in Deinotherium (largely based on Harris, 1975), Gomphotherium (largely based on Tassy, 2013), and Loxodonta (based on Boas & Paulli, 1908 and Eales, 1926), with arrows showing directionality of m. temporalis and m. masseter muscles. (b) Phylogeny of proboscideans (based on studies referenced by Sanders, 2023) showing muscles of mastication and hypothetical proboscis morphology, with directionality of feeding mechanisms labeled (not including integral orthal feeding motions). Not to scale.
Of tusks and trunks: A review of craniofacial evolutionary anatomy in elephants and extinct Proboscidea

October 2024

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414 Reads

While being the largest living terrestrial mammals, elephants are best known for their highly modified and uniquely elaborate craniofacial anatomy—most notably with respect to their often‐massive tusks and intricately muscular, multifunctional proboscis (i.e., trunk). For over a century, studies of extinct proboscidean relatives of today's elephants have presented hypotheses regarding the evolutionary history of the crania and tusks of these animals and their bearing on the evolution of the proboscis. Herein, I explore major functional characteristics of the proboscidean head. I give a brief review of the anatomy of tusks and dentition, the feeding apparatus, and proboscis in extant elephants and explore their overall bearing in elephant feeding behavior as well as other aspects of their ecology. I also review the evolution of the proboscidean head, with a synthetic analysis of studies and further speculation exploring the interconnected evolutionary roles of tusk morphology and use, feeding anatomy and functional implications thereof, and proboscis anatomy and use in the ancestry of elephants. Notable emphasis is given to the evolutionary role of initial elongation of the mandibular symphysis in the development of the proboscis in many proboscideans. Subsequent secondary shortening of the symphysis and elevation of the temporal region and occiput allowed for a pendulous trunk and proal feeding in living elephants and other proboscidean groups with highly lophodont dentition.


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1.8 (2023)

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45%

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4.8 (2023)

CiteScore™


16 days

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$4,220 / £2,870 / €3,520

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