Preliminary ontogeny of the giraffe neck.

The giraffe juvenile has different proportions of head to neck from the adult. The head just about doubles in size from the juvenile to adult, whereas the neck increases almost 4.5× (roughly four times) in length. The T1 posterior dorsal vertebral width of the newborn is clearly wider than in the adult where it is narrow. In the okapi, the dorsal vertebral width is narrow in both juvenile and adult. The giraffe neck changes in ontogeny anisometrically. In the okapi the changes are more isometric. The giraffe juvenile vertebrae are shorter and do not have fused the cranial epiphyseal plates. That facilitates anterior elongation-growth. The ventral tubercles are undeveloped. The juvenile T1 is wide caudally unlike the adult. This may be a similarity to a gelocid (Gelocidae) ancestor of the giraffe.

Comment on "Sexual selection promotes giraffoid head-neck evolution and ecological adaptation".

Wang et al. (Research Articles, 3 June 2022, eabl8316) reported an early Miocene giraffoid that exhibited fierce head-butting behavior and concluded that sexual selection promoted head-neck evolution in giraffoids. However, we argue that this ruminant is not a giraffoid and thus that the hypothesis that sexual selection promoted giraffoid head-neck evolution is not sufficiently supported.

The bony cap and its distinction from the distal phalanx in humans, cats, and horses.

It has been recognized as early as the Victorian era that the apex of the distal phalanx has a distinct embryological development from the main shaft of the distal phalanx. Recent studies in regenerative medicine have placed an emphasis on the role of the apex of the distal phalanx in bone regrowth. Despite knowledge about the unique aspects of the distal phalanx, all phalanges are often treated as equivalent. Our morphological study reiterates and highlights the special anatomical and embryological properties of the apex of the distal phalanx, and names the apex "the bony cap" to distinguish it. We posit that the distal phalanx shaft is endochondral, while the bony cap is intramembranous and derived from the ectodermal wall. During development, the bony cap may be a separate structure that will fuse to the endochondral distal phalanx in the adult, as it ossifies well before the distal phalanges across taxa. Our study describes and revives the identity of the bony cap, and we identify it in three mammalian species: humans, cats, and horses (Homo sapiens, Felis catus domestica, and Equus caballus). During the embryonic period, we show the bony cap has a thimble-like shape that surrounds the proximal endochondral distal phalanx. The bony cap may thus play an inductive role in the differentiation of the corresponding nail, claw, or hoof (keratin structures) of the digit. When it is not present or develops erroneously, the corresponding keratin structures are affected, and regeneration is inhibited. By terming the bony cap, we hope to inspire more attention to its distinct identity and role in regeneration.

Comparisons of Schansitherium tafeli with Samotherium boissieri (Giraffidae, Mammalia) from the Late Miocene of Gansu Province, China.

We are describing and figuring for the first time skulls of Schansitherium tafeli, which are abundant in the Gansu area of China from the Late Miocene. They were animals about the size of Samotherium with shorter necks that had two pairs of ossicones that merge at the base, which is unlike Samotherium. The anterior ossicones consist of anterior lineations, which may represent growth lines. They were likely mixed feeders similar to Samotherium. Schansitherium is tentatively placed in a very close position to Samotherium. Samotherium and Schansitherium represent a pair of morphologically very similar species that likely coexisted similarly to pairs of modern species, where the main difference is in the ossicones. Pairs of ruminants in Africa, for example, exist today that differ mostly in their horn shape but otherwise are similar in size, shape, and diet. The absence of Schansitherium from Europe is interesting, however, as Samotherium is found in both locations. While is it challenging to interpret neck length and ossicone shape in terms of function in combat, we offer our hypothesis as to how the two species differed in their fighting techniques.

The evolution and anatomy of the horse manus with an emphasis on digit reduction.

We revisit digit reduction in the horse and propose that all five digits are partially present in the modern adult forelimb. Osteological descriptions of selected tetradactyl, tridactyl and monodactyl equids demonstrate the evolution of the forelimb. Histological, osteological and palaeontological evidence suggest that the Equus distal forelimb is more complex than traditionally conceived. The current understanding is that the horse distal forelimb consists of one complete digit (III) and two reduced splint metacarpals (II and IV). Metacarpals II and IV each exhibit a ventral ridge, which we suggest represents the undifferentiated digits I and V. These ridges are present in the tridactyl Mesohippus, but are absent in the tetradactyl Hyracotherium. The carpal articulations of the five metacarpals match those of pentadactyl taxa. Distally, the frog, a V-shaped structure on the ventral hoof represents digits II and IV, and the wings and hoof cartilages of the distal phalanx are digits I and V. We relate this revised interpretation of the Equus forelimb to Laetoli footprints, and suggest the Hipparion side impressions are created from the hooves of I and V, rather than from II and IV. We show shades of pentadactyly within the Equus manus.

The earliest ossicone and post-cranial record of Giraffa.

The oldest Giraffa material presently known consists of dental specimens. The oldest post-cranial Giraffa material belongs to the Plio-Pleistocene taxon Giraffa sivalensis, where the holotype is a third cervical vertebra. We describe three non-dental specimens from the Early Late Miocene of the Potwar Plateau, including an 8.1 million year old ossicone, 9.4 million year old astragalus, and 8.9 million year old metatarsal and refer them to Giraffa. The described ossicone exhibits remarkable similarities with the ossicones of a juvenile modern giraffe, including the distribution of secondary bone growth, posterior curvature, and concave pitted undersurface where the ossicone would attach to the skull. The astragalus has a notably flat grove of the trochlea, medial twisting between the trochlea and the head, and a square-shaped sustentacular facet, all of which characterize the astragalus of Giraffa camelopardalis. The newly described astragalus is narrow and rectangular, unlike the boxy shaped bone of the modern giraffe. The metatarsal is large in size and has a shallow central trough created by thin medial and lateral ridges, a feature unique to Giraffa and Sivatherium. Our described material introduce the earliest non-dental material of Giraffa, a genus whose extinct representation is otherwise dominated by teeth, and demonstrate that the genus has been morphologically consistent over 9 million years.

Postembryonic Nephrogenesis and Persistence of Six2-Expressing Nephron Progenitor Cells in the Reptilian Kidney.

New nephron formation (nephrogenesis) ceases in mammals around birth and is completely absent in adults. In contrast, postembryonic nephrogenesis is well documented in the mesonephric kidneys of fishes and amphibians. The transient mesonephros in reptiles (including birds) and mammals is replaced by the metanephros during embryogenesis. Thus, one may speculate that postembryonic nephrogenesis is restricted to the mesonephric kidney. Previous reports have suggested the metanephros of non-avian reptiles (hereafter reptiles) may continually form nephrons throughout life. We investigated the presence of adult nephrogenesis in reptiles by examining adult kidneys from several species including Trachemys scripta, Chrysemys picta, Boa constrictor, Tupinambis tegu, Anolis carolinensis, and Alligator mississipiensis among others. We found that all major reptilian groups (Testudines, Crocodylia, and Squamates) showed the presence of adult nephrogenesis. The total amount of nephrogenesis varied greatly between species with turtles displaying the highest density of nephrogenesis. In contrast, we were unable to detect adult nephrogenesis in monotremes, and in the iguanid A. carolinensis. Nephron progenitor cells express the transcription factor Six2, which in mammals, becomes downregulated as the progenitor cell population is exhausted and nephrogenesis ends. Using the alligator as a model, we were able to detect Six2-positive cap mesenchyme cells in the adult kidney, which spatially correlated with areas of nephrogenesis. These results suggest that the metanephric kidney of reptiles has maintained the ability to continually grow new nephrons during postembryonic life, a process lost early in mammalian evolution, likely due to the persistence of a Six2-expressing progenitor cell population.

Astragalar Morphology of Selected Giraffidae.

The artiodactyl astragalus has been modified to exhibit two trochleae, creating a double pullied structure allowing for significant dorso-plantar motion, and limited mediolateral motion. The astragalus structure is partly influenced by environmental substrates, and correspondingly, morphometric studies can yield paleohabitat information. The present study establishes terminology and describes detailed morphological features on giraffid astragali. Each giraffid astragalus exhibits a unique combination of anatomical characteristics. The giraffid astragalar morphologies reinforce previously established phylogenetic relationships. We find that the enlargement of the navicular head is a feature shared by all giraffids, and that the primitive giraffids possess exceptionally tall astragalar heads in relation to the total astragalar height. The sivatheres and the okapi share a reduced notch on the lateral edge of the astragalus. We find that Samotherium is more primitive in astragalar morphologies than Palaeotragus, which is reinforced by tooth characteristics and ossicone position. Diagnostic anatomical characters on the astragalus allow for giraffid species identifications and a better understanding of Giraffidae.

The cervical anatomy of Samotherium, an intermediate-necked giraffid.

Giraffidae are represented by many extinct species. The only two extant taxa possess diametrically contrasting cervical morphology, as the okapi is short-necked and the giraffe is exceptionally long-necked. Samotherium major, known from the Late Miocene of Samos in Greece and other Eurasian localities, is a key extinct giraffid; it possesses cervical vertebrae that are intermediate in the evolutionary elongation of the neck. We describe detailed anatomical features of the cervicals of S. major, and compare these characteristics with the vertebrae of the two extant giraffid taxa. Based on qualitative morphological characters and a quantitative analysis of cervical dimensions, we find that the S. major neck is intermediate between that of the okapi and the giraffe. Specifically, the more cranial (C2-C3) vertebrae of S. major represent a mosaic of features shared either with the giraffe or with the okapi. The more caudal (C5-C7) S. major vertebrae, however, appear transitional between the two extant taxa, and hence are more unique. Notably, the C6 of S. major exhibits a partially excavated ventral lamina that is strong cranially but completely absent on the caudal half of the ventral vertebral body, features between those seen in the giraffe and the okapi. Comprehensive anatomical descriptions and measurements of the almost-complete cervical column reveal that S. major is a truly intermediate-necked giraffid. Reconstructions of the neck display our findings.

Fossil evidence and stages of elongation of the Giraffa camelopardalis neck.

Several evolutionary theories have been proposed to explain the adaptation of the long giraffe neck; however, few studies examine the fossil cervical vertebrae. We incorporate extinct giraffids, and the okapi and giraffe cervical vertebral specimens in a comprehensive analysis of the anatomy and elongation of the neck. We establish and evaluate 20 character states that relate to general, cranial and caudal vertebral lengthening, and calculate a length-to-width ratio to measure the relative slenderness of the vertebrae. Our sample includes cervical vertebrae (n=71) of 11 taxa representing all seven subfamilies. We also perform a computational comparison of the C3 of Samotherium and Giraffa camelopardalis, which demonstrates that cervical elongation occurs disproportionately along the cranial-caudal vertebral axis. Using the morphological characters and calculated ratios, we propose stages in cervical lengthening, which are supported by the mathematical transformations using fossil and extant specimens. We find that cervical elongation is anisometric and unexpectedly precedes Giraffidae. Within the family, cranial vertebral elongation is the first lengthening stage observed followed by caudal vertebral elongation, which accounts for the extremely long neck of the giraffe.

The Cervical Osteology of Okapia johnstoni and Giraffa camelopardalis.

Giraffidae is the only family of ruminants that is represented by two extant species; Okapia johnstoni and Giraffa camelopardalis. Of these taxa, O. johnstoni represents a typical short-necked ungulate, and G. camelopardalis exemplifies the most extreme cervical elongation seen in any ruminant. We utilize these two species to provide a comprehensive anatomic description of the cervical vertebrae. In addition, we compare the serial morphologic characteristics of the okapi and giraffe cervical vertebrae, and report on several osteologic differences seen between the two taxa. The giraffe neck appears to exhibit homogenization of C3-C7; the position of the dorsal tubercle, thickness of the cranial articular process, shape of the ventral vertebral body, and orientation of the ventral tubercle are constant throughout these vertebrae, whereas these features are serially variable in the okapi. We also report on several specializations of the giraffe C7, which we believe relates to an atypical cervico-thoracic junction, corresponding to the substantial neck lengthening. The morphologic differences exhibited between the okapi and giraffe cervical vertebrae have implications on the function of the necks relating to both fighting and feeding.

Posterior subscapular dissection: An improved approach to the brachial plexus for human anatomy students.

Students of human anatomy are required to understand the brachial plexus, from the proximal roots extending from spinal nerves C5 through T1, to the distal-most branches that innervate the shoulder and upper limb. However, in human cadaver dissection labs, students are often instructed to dissect the brachial plexus using an antero-axillary approach that incompletely exposes the brachial plexus. This approach readily exposes the distal segments of the brachial plexus but exposure of proximal and posterior segments require extensive dissection of neck and shoulder structures. Therefore, the proximal and posterior segments of the brachial plexus, including the roots, trunks, divisions, posterior cord and proximally branching peripheral nerves often remain unobserved during study of the cadaveric shoulder and brachial plexus. Here we introduce a subscapular approach that exposes the entire brachial plexus, with minimal amount of dissection or destruction of surrounding structures. Lateral retraction of the scapula reveals the entire length of the brachial plexus in the subscapular space, exposing the brachial plexus roots and other proximal segments. Combining the subscapular approach with the traditional antero-axillary approach allows students to observe the cadaveric brachial plexus in its entirety. Exposure of the brachial dissection in the subscapular space requires little time and is easily incorporated into a preexisting anatomy lab curriculum without scheduling additional time for dissection.

Dietary change and evolution of horses in North America.

The evolution of high-crowned molars among horses (Family Equidae) is thought to be an adaptation for abrasive diets associated with the spread of grasslands. The sharpness and relief of the worn cusp apices of teeth (mesowear) are a measure of dietary abrasion. We collected mesowear data for North American Equidae for the past 55.5 million years to test the association of molar height and dietary abrasion. Mesowear trends in horses are reflective of global cooling and associated vegetation changes. There is a strong correlation between mesowear and crown height in horses; however, most horse paleopopulations had highly variable amounts of dietary abrasion, suggesting that selective pressures for crown height may have been weak much of the time. However, instances of higher abrasion were observed in some paleopopulations, suggesting intervals of stronger selection for the evolution of dentitions, including the early Miocene shortly before the first appearance of Equinae, the horse subfamily in which high-crowned dentitions evolved.

Can low-magnification stereomicroscopy reveal diet?

A new method of scoring dental microscopic use wear, initially developed for and applied to extant and extinct ungulates, is here applied to primates, and the efficacy of the method as a tool for diagnosing diet in both ungulates and primates is established. The method employs standard refractive light microscopy instead of scanning electron microscopy (SEM), and all use-wear features are counted or scored under low magnification (35 x). We use measurement systems analysis (variance components analysis of sources of measurement error) to evaluate the consistency and reproducibility of measurements using this method. The method is shown to have low intra- and inter-observer measurement error, and to effectively distinguish among graminivores, folivores, and frugivores. It can also be used to identify seed predators and to diagnose hard-object feeding. The method is also shown to be robust to the selection of measurement site; it works equally well when applied to upper or to lower molars. Finally, we use analysis of variance to examine the consistency of the signals across mammalian orders, and discriminant function analysis to develop dietary diagnostic tools for a set of "classified" primates with known diets. We test the success of these tools not merely by examining their a posteriori classification "success," but by using them to construct predicted dietary profiles for a sample of unclassified extant primate species, again with known diets.

Dental use wear in extinct lemurs: evidence of diet and niche differentiation.

A new technique for molar use-wear analysis is applied to samples of all 16 species of extinct lemurs with known dentitions, as well as to a large comparative sample of extant primates. This technique, which relies on the light refractive properties of wear pits and scratches as seen under a standard stereoscopic microscope, has shown itself to be effective in distinguishing the diets of ungulates and extant primates. We draw dietary inferences for each of the 16 extinct lemur species in our database. There is a strong phylogenetic signal, with the Palaeopropithecidae showing use-wear signatures similar to those of the Indriidae; extinct lemurids (Pachylemur spp.) showing striking similarities to extant lemurids (except Hapalemur spp.); and Megaladapis showing similarities to Lepilemur spp. Only the Archaeolemuridae have dietary signatures unlike those of any extant lemurs, with the partial exception of Daubentonia. We conclude that the Archaeolemuridae were hard-object feeders; the Palaeopropithecidae were seed predators, consuming a mixed diet of foliage and fruit to varying degrees; Pachylemur was a fruit-dominated mixed feeder, but not a seed predator; and all Megaladapis were leaf browsers. There is no molar use wear evidence that any of the extinct lemurs relied on terrestrial foods (C4 grasses, tubers, rhizomes). This has possible implications for the role of the disappearance of wooded habitats in the extinction of lemurs.

Approach to the educational opportunities provided by variant anatomy, illustrated by discussion of a duplicated inferior vena cava.

Variant anatomy recognized during routine cadaveric dissection in the first year of medical school offers great learning potential by allowing students to gain enhanced understanding of an array of important subjects. It provides a framework for reviewing common morphology and embryogenesis of the structure in question, and through the help of appropriate faculty, yields insight into the potential medical, radiologic, and surgical implications. The frequency of clinically important anatomic variation is high enough to allow the gross anatomy laboratory to serve as an excellent teaching platform in this regard. Through anatomy, the student is introduced to the concept of patient individuality, and to the individualization of medical and surgical therapies. Recently, one of the variations encountered in our lab was a duplicated inferior vena cava. We describe our approach to such findings through a systematic discussion of the anatomy and embryology, as well as the radiologic and clinical correlates.