Equid Nutritional Physiology and Behavior: An Evolutionary Perspective.

Like other members of the odd-toed ungulates (the perissodactyls), equids once had a higher species diversity in the fossil record than they have today. This is generally explained in comparison to the enormous diversity of bovid ruminants. Theories on putative competitive disadvantages of equids include the use of a single toe as opposed to two toes per leg, the lack of a specific brain cooling (and hence water-saving) mechanism, longer gestation periods that delay reproductive output, and in particular digestive physiology. To date, there is no empirical support for the theory that equids fare better on low-quality forage than ruminants. In contrast to the traditional juxtaposition of hindgut and foregut fermenters, we suggest that it is more insightful to sketch the evolution of equid and ruminant digestive physiology as a case of convergence: both evolved a particularly high chewing efficacy in their respective groups, which facilitates comparatively high feed and hence energy intakes. But because the ruminant system, less based on tooth anatomy but more on a forestomach sorting mechanism, is more effective, equids depend more on high feed intakes than ruminants and may well be more susceptible to feed shortages. Arguably, the most underemphasized characteristic of equids may be that in contrast to many other herbivores including ruminants and coprophageous hindgut fermenters, equids do not use the microbial biomass growing in their gastrointestinal tract. Equids display behavioral and morphophysiological adaptations to high feed intakes, and their cranial anatomy that facilitates the cropping of forage while performing grinding chewing at the same time might be unique. Rather than looking for explanations how equids are better adapted to their present niches than other organisms, considering them remnants of a different morphophysiological solution may be more appropriate.

Silicone plastination of nasus and lingua in large ruminants with and without use of formaldehyde / Plastinación de silicona de nasus y lengua en rumiantes mayores con y sin el uso de formaldehido

SUMMARY: Plastination is an anatomical preparate preparation technique characterized by the replacement of tissue fluids with a reactive polymer. Although more challenging and economically costly than many anatomical methods, this method is desirable because of the fact that specimens created in this method are highly similar to the natural appearance of the intended objects, and they are durable and harmless end products for human health. Our main goal was to completely leave out formaldehyde and similar carcinogenic chemicals used in a method like plastination and to allow production of formaldehyde-free plastinates to be used in anatomy training and examinations in our country. To that end, we compared nose and tongue of 10 large ruminants by subjecting them to plastination, 5 of them with formaldehyde and 5 of them without formaldehyde, and aimed to leave formaldehyde out by taking into account the difference between them. Silicone plastination is the most commonly-used and best-known technique among the plastination techniques because specimens created using this technique look aesthetically impressive. Silicone plastination consists mainly of 5 phases. First of all, we obtained the anatomical situs we wanted and made specimens ready by dissecting some of them after fixation and some of them without fixation. Then, after the implementation of a dehydration phase in acetone baths at -25 °C, a forced impregnation phase was implemented by using a mixture of S10-S3 chemical under negative pressure. In the final phase, the curing and hardening phase, the plastination process was completed by giving the specimens their final shape with the use of the S6 solution. As a result, no significant difference was observed between silicone plastination with and without formaldehyde.

RESUMEN: La plastinación es una técnica de preparados anatómicos caracterizada por la sustitución de fluidos tisulares por un polímero reactivo. A pesar de ser económicamente más costoso que muchas métodos anatómicos, este técnica es deseable debido a que las muestras creadas son muy similares a la apariencia natural de los objetos previstos y son productos finales duraderos e inofensivos para la salud humana. Nuestro objetivo principal fue dejar completamente de lado el formaldehído y las sustancias químicas cancerígenas similares utilizadas en un método como la plastinación y permitir la producción de plastinados libres de formaldehído para su uso en la formación y los exámenes de anatomía en nuestro país. Con ese fin, comparamos la nariz y la lengua de 10 rumiantes mayores sometiéndolos a plastinación, 5 de ellos con formaldehído y 5 de ellos sin formaldehído, y buscamos eliminar el formaldehído considerando la diferencia entre ellos. La plastinación con silicona es la técnica más utilizada y más conocida entre las técnicas de plastinación porque los especímenes creados con ella se ven estéticamente impresionantes. La plastinación con silicona consta principalmente de 5 fases. En primer lugar, obtuvimos el situs anatómico que queríamos y preparamos los especímenes diseccionando algunos de ellos después de la fijación y otros sin fijación. Luego, de la implementación de una fase de deshidratación en baños de acetona a -25 °C, se implementó una fase de impregnación forzada utilizando una mezcla del químico S10-S3 a presión negativa. En la fase final, la fase de curado y endurecimiento, se completó el proceso de plastinación dando a los especímenes su forma definitiva con el uso de la solución S6. Como resultado, no se observaron diferencias significativas entre la plastinación con silicona con y sin formaldehído.

Histological architectures and biometric characteristics of indigenously plastinated organs of goat / Arquitectura histológica y características biométricas de órganos de cabra plastinados localmente

SUMMARY: The histo-morphology and biometry of liver, lung and kidney of goat were studied. The obtained samples were divided into two groups. One set of tissue was processed for paraffin embedding after formalin fixation. The other set was indigenously plastinated. The plastinates were embedded with paraffin without deplastination. Both the non plastinated and plastinated tissues were sectioned and processed for routine staining. The tissues were examined under light microscope for histological architectures and quantitatively assessed the biometric parameters. The non plastinated and plastinated samples yielded mostly similar histological architectures. But plastinated liver showed alterations and artifacts with enlargement of the central vein and hepatic sinusoidal space. Plastinated lung revealed enlarged alveolar sac. Lack of nuclear clarity was observed for all the plastinated samples. The plastinated kidney revealed compactness of the cellular structures and shrinkage induced artifacts with clear renal corpuscles structure and obvious glomerular capsule (Bowman's capsule). The biometric measurements of central vein of liver, alveolar sac of lung, glomerulus and glomerular space of kidney of both plastinated and non plastinated tissue were also performed using calibrated stage micrometer. Comparison was done between the biometric data for both plastinated and non plastinated tissue. No significant difference was found in the obtained biometric data of liver and lung. Only a significant difference was observed between the width of glomerulus of non plastinated kidney and width of glomerulus of plastinated kidney. The morphological and biometric study of non plastinated and plastinated organs of goat can conclude that histological slides can be made from indigenously plastinated organs with well preserved histological architectures of the tissues with some rare exceptions. Plastination does not affect the biometric characteristics of the organs. In addition, the comparative morphologic and biometric study of plastinated and non plastinated organs of goat will be useful in education and research.

RESUMEN: En este estudio se analizó la histomorfología y biometría de hígado, pulmón y riñón de cabra. Las muestras obtenidas se dividieron en dos grupos: una cohorte de tejido fue procesada para su inclusión en parafina después de la fijación con formalina. La siguiente cohorte fue plastinada localmente. Tanto los tejidos no plastinados como los plastinados fueron seccionados y procesados para tinción de rutina. Los tejidos se examinaron con un microscopio óptico en busca de arquitecturas histológicas y se evaluaron cuantitativamente los parámetros biométricos. Las muestras no plastinadas y plastinadas produjeron arquitecturas histológicas en su mayoría similares. El hígado plastinado mostró alteraciones y artefactos con un aumento de la vena central y del espacio sinusoidal hepático. El pulmón plastinado reveló aumento del saco alveolar. Se observó falta de claridad nuclear en todas las muestras plastinadas. El riñón plastinado reveló compacidad de las estructuras celulares y artefactos inducidos por contracción con estructura clara de corpúsculos renales y obvia cápsula glomerular (cápsula de Bowman). Las mediciones biométricas de la vena central del hígado, el saco alveolar del pulmón, el glomérulo y el espacio glomerular del riñón de tejido plastinado y no plastinado también se realizaron utilizando un micrómetro de platina calibrado. Se realizó una comparación entre los datos biométricos del tejido plastinado y no plastinado. No se encontraron diferencias significativas en los datos biométricos obtenidos de hígado y pulmón. Se observó una diferencia significativa entre el ancho del glomérulo del riñón no plastinado y el ancho del glomérulo del riñón plastinado. En el estudio morfológico y biométrico de órganos de cabra no plastinados y plastinados se puede concluir que es posible hacer portaobjetos histológicos a partir de órganos plastinados naturales con arquitecturas histológicas de los tejidos bien conservadas, con algunas excepciones. La plastinación no afecta las características biométricas de los órganos. Además, el estudio comparativo morfológico y biométrico de órganos plastinados y no plastinados de cabra será de utilidad en la educación y la investigación.

Anatomy of the skull of saanen goat: an anesthesiology and stereology approach / Anatomía del cráneo de la cabra saanen: un enfoque desde la anestesiología y la estereología

SUMMARY: The Saanen goat is known as the greatest milk producer among small ruminat breeds. However, its morphometric features still remain unclear. Therefore, the present work aimed to investigate the functional anatomy of the upper and lower jaws as well as the volumetric properties of the male and female Saanen goat for clinical applications. The heads of 20 adult animals (10 males and 10 females) were included. Totally, 22 morphometric parameters were measured on three dimensional computed tomographic images using RadiAnt DICOM Viewer software and some parameters were measured on hot macerated sample. The mean volume of paranasal sinuses as well as conchal sinuses were estimated using stereological method. Based on the results, the differences between males and females were not significant (p>0.05) in all desired parameters.The frontal and lacrimal sinuses were the largest and smallest paranasal sinus in both sexes. Also, the dorsal and middle conchal sinuses were the largest and smallest ones, respectively. The common nasal meatus was the smallest and ventral meatus was the largest meatus in the nasal cavity. In conclusion, these findings provide a basic data that would be useful in blocking terminal branches of the cranial nerves in this breed for surgical purpose or teeth injuries treatment.

RESUMEN: La cabra Saanen es conocida como la mayor productora de leche entre las razas de pequeños ruminos, sin embargo, sus características morfométricas aún permanecen sin revelar. Por lo tanto, el presente trabajo tuvo como objetivo investigar la anatomía de los huesos del cráneo y cara, así como sus propiedades volumétricas, en la cabra Saanen, tanto del macho como de la hembra con la finalidad de contribuir con las aplicaciones clínicas. Se incluyeron las cabezas de 20 animales adultos (10 machos y 10 hembras). Se midieron 22 parámetros morfométricos en imágenes de tomografía computarizada tridimensionales utilizando el software RadiAnt DICOM Viewer; algunos parámetros se midieron en muestra macerada en calor. El volumen medio de los senos paranasales y concales se estimó mediante método estereológico. En base a los resultados, las diferencias entre machos y hembras no fueron significativas (p> 0.05) en todos los parámetros deseados. Los senos frontal y lagrimal eran de mayor y menor volumen en ambos sexos, respectivamente. Además, los senos conchal dorsal y medio eran los más grandes y los más pequeños, respectivamente. El meato nasal común fue el más pequeño y el meato ventral el más grande en la cavidad nasal. En conclusión, estos hallazgos brindan un dato básico que sería útil en el bloqueo de las ramas terminales de los nervios craneales en esta raza, con fines quirúrgicos o tratamiento de lesiones dentales.

Reassessment of the enigmatic ruminant Miocene genus Amphimoschus Bourgeois, 1873 (Mammalia, Artiodactyla, Pecora).

Amphimoschus is an extinct Eurasian ruminant genus, mostly recorded in Europe, without a close living relative and, hence, an unknown systematic position. This genus is known from around 50 localities from the late early to the middle Miocene. Two species were described during 180 years, but since their first description during the late 19th century and early 20th century, hardly any detailed taxonomic work has been done on the genus. Over the years, extensive collecting and excavating activities have enriched collections with more and more complete material of this still rare and enigmatic animal. Most interestingly, a number of skull remains have been unearthed and are promising in terms of providing phylogenetic information. In the present paper, we describe cranial material, the bony labyrinth, the dentition through 780 teeth and five skulls from different ontogenetic stages. We cannot find a clear morphometric distinction between the supposedly smaller and older species Amphimoschus artenensis and the supposedly younger and larger species A. ponteleviensis. Accordingly, we have no reason to retain the two species and propose, following the principle of priority (ICZN chapter 6 article 23), that only A. ponteleviensis Bourgeois, 1873 is valid. Our studies on the ontogenetic variation of Amphimoschus does reveal that the sagittal crest may increase in size and a supraorbital ridge may appear with age. Despite the abundant material, the family affiliation is still uncertain.

Imaging of the Urinary and Reproductive Tract in Small Ruminants.

Ultrasound imaging is the mainstay of urinary and reproductive imaging in small ruminants, assisted by the relatively superficial location of the urinary and reproductive tracts in these species. Radiography can provide an excellent overview of the abdomen, but is often limited by a lack of adequate penetration. Computed tomography scans provide summation-free imaging of the urinary and reproductive tracts in addition to information about the excretion of contrast agent by the kidneys, but can be limited by availability and cost.

Formation, structure, and function of extra-skeletal bones in mammals.

This review describes the formation, structure, and function of bony compartments in antlers, horns, ossicones, osteoderm and the os penis/os clitoris (collectively referred to herein as AHOOO structures) in extant mammals. AHOOOs are extra-skeletal bones that originate from subcutaneous (dermal) tissues in a wide variety of mammals, and this review elaborates on the co-development of the bone and skin in these structures. During foetal stages, primordial cells for the bony compartments arise in subcutaneous tissues. The epithelial-mesenchymal transition is assumed to play a key role in the differentiation of bone, cartilage, skin and other tissues in AHOOO structures. AHOOO ossification takes place after skeletal bone formation, and may depend on sexual maturity. Skin keratinization occurs in tandem with ossification and may be under the control of androgens. Both endochondral and intramembranous ossification participate in bony compartment formation. There is variation in gradients of density in different AHOOO structures. These gradients, which vary according to function and species, primarily reduce mechanical stress. Anchorage of AHOOOs to their surrounding tissues fortifies these structures and is accomplished by bone-bone fusion and Sharpey fibres. The presence of the integument is essential for the protection and function of the bony compartments. Three major functions can be attributed to AHOOOs: mechanical, visual, and thermoregulatory. This review provides the first extensive comparative description of the skeletal and integumentary systems of AHOOOs in a variety of mammals.

The relative importance of phylogeny and habitat in determining the presence and prominence of a granula iridica in hooved mammals.

PURPOSE: To investigate the relationship between phylogeny and amount of shade in a species' habitat regarding the presence or absence of an iridal granula iridica (GI) in a large sample of Artiodactyl and Perissodactyl clades and using online resources. METHODS: The Comparative Ocular Pathology Laboratory of Wisconsin (COPLOW) archives were searched for glass slide material from Artiodactyl (even-toed) and Perissodactyl (odd-toed) ungulates. The slides were examined, and the presence or absence of the GI was noted. The phylogenetic tree of the ungulate species was inferred using TimeTree (http://www.timetree.org), and the habitat data are derived from Animal Diversity Web (https://animaldiversity.org/). We assessed the probability of the presence of GI occurring given the amount of shade in a species' environment using phylogenetic logistic regression. RESULTS: Forty-eight artiodactyl species were able to be evaluated and tabulated. Nine perissodactyl species were able to be evaluated. The phylogenetic logistic regression showed that the probability of GI presence was lower in artiodactyl species that inhabited shaded environments (ßshaded  = -1.774). Arctiodacyl species inhabiting a nonshaded environment were slightly more probable to have the GI present (ßnonshaded  = 0.023), with species inhabitating ambiguously shaded environments having a high probability of GI presence (ßambiguous  = 2.214). CONCLUSIONS: Our results suggest that the GI may be a common morphological feature to shade the pupil in nonshaded environments, and, in its absence, increase the amount of light reaching the retina to improve vision in shaded environments for hooved mammals. Further research on the functional optics of the GI and studies that include additional ungulate species would further elucidate phylogenetic and ecological factors influencing the occurrence of GI in hooved mammals.

Comparative study of the eyelids and orbital glands morphology in the okapi (Okapia johnstoni, Giraffidae), Père David's deer (Elaphurus davidianus, Cervidae) and the Philippine mouse-deer (Tragulus nigricans, Tragulidae).

The accessory organs of the eye represent part of the protective system of the eyeball. In the present study, an examination of the accessory organs of the eye of three species of captive ruminants was performed using light microscopy. In the okapi, the superficial gland of the third eyelid and lacrimal gland were complex branched multilobar tubular glands formed by mucous units with tubular secretory portions and no plasma cells. The deep gland of the third eyelid was absent in the okapi and present in both the Père David's deer and the Philippine mouse-deer. In the Philippine mouse-deer, the deep gland had a very thick connective capsule and thick interlobar septae. It contained fewer lobes forming the gland parenchyma compared to Père David's deer and other ruminants. Organized lymphoid follicles were present within the upper and lower eyelids only in the okapi and Père David's deer, while diffuse lymphocytes were observed in the Philippine mouse-deer. The orbital glands in the Père David's deer had a multilobar tubuloacinar structure with numerous plasma cells and a mucoserous character. In contrast to the Philippine mouse-deer, these glands had a serous character. The presence of several macroscopic and microscopic structural differences of the examined accessory organs of the eye in the three captive ruminant species may be understood within an ecological context and may be associated with different habitat-specific environmental conditions.

Ruminants (Giraffidae and Bovidae) from Kanapoi.

We update here our recent revision of the Kanapoi ruminants and describe recently collected material. We now regard the occurrence of reduncins as doubtful, we revise the identification of a large raphicerin as being more probably Gazella, and we add Gazella cf. janenschi and the Cephalophini to the faunal list. New material of Tragelaphus kyaloi suggests that this species held its head unlike other tragelaphins, and was not an exclusive dedicated browser, but Kanapoi pre-dates the Pliocene change of Sivatherium, Aepyceros, Alcelaphini, and even Tragelaphini toward more grazing diets. Kanapoi shares several ruminant taxa with sites in Ethiopia and Tanzania, attesting to latitudinal exchanges.

Intraspecific male combat behaviour predicts morphology of cervical vertebrae in ruminant mammals.

Cranial weapons of all shapes and sizes are common throughout the animal kingdom and are frequently accompanied by the evolution of additional traits that enhance the use of those weapons. Bovids (cattle, sheep, goats, antelope) and cervids (deer) within the mammal clade Ruminantia are particularly well known for their distinct and varied cranial appendages in the form of horns and antlers, which are used as weapons in intraspecific combat between males for access to mates. Combat in these species takes many forms, including head-on collisions (ramming); stabbing an opponent's head or body with horn tips (stabbing); rearing and clashing downwards with horns (fencing); or interlocking antlers or horns while vigorously pushing and twisting (wrestling). Some aspects of weapon and skull morphology have been linked to combat behaviours in bovid and cervid species, but the contribution of postcranial structures that support these weapons, such as the neck, has not been explored. To investigate the role of the neck in intraspecific combat, we quantified biomechanically relevant linear variables of the cervical vertebrae (C1-C7) from males and females of 55 ruminant species. We then used phylogenetic generalized least-squares regression to assess differences among species that display primarily ramming, stabbing, fencing and wrestling combat styles. In males, we found that wrestlers have longer vertebral centra and longer neural spines than rammers, stabbers or fencers, while rammers have shorter and wider centra and taller neural spine lever arms. These results suggest a supportive role for the cervical vertebrae in resisting forces generated by male-male combat in ruminant mammals and indicate that evolutionary forces influencing cranial weapons also play a role in shaping the supporting anatomical structures.

Morphology of the lingual papillae of the bharal (Pseudois nayaur).

We examined the dorsal lingual surfaces of an adult bharal (Pseudois nayaur) by scanning electron microscopy. The filiform papillae of the lingual apex and body consisted of a main papilla and smaller secondary papillae. The filiform papilla of the lingual body was big as compared to that of the lingual apex. The connective tissue cores of the filiform papillae consisted of several processes. The fungiform papilla was round in shape. The connective tissue cores of the fungiform papillae were flower-bud shaped. The lenticular papillae of large size were limited on the lingual prominence. The connective tissue cores of the lenticular papillae were hair-like in shape. The vallate papillae were located on both sides of the posterolateral aspects. The vallate papillae were flattened-oval shaped and the papillae were surrounded by an oval-shaped trench. The connective tissue cores of the vallate papillae were covered with numerous small spines The lingual surface of the bharal closely resembled that of the family Bovidae.

Gross intestinal morphometry and allometry in ruminants.

While some descriptions of ruminants' dietary adaptations suggest that the length of the intestinal tract reflects the proportion of grass or browse in the diet, this assumption has been questioned. We collated data on body mass (BM), as well as small intestine, caecum, colon/rectum, large and total intestine length in 68 ruminant species, and, while accounting for the phylogenetic structure of the dataset, evaluated both allometric scaling and the potential influence of diet, digestive physiology or climate proxies on measures of intestine length. Intestinal length generally scaled to BM at an exponent higher than the 0.33 expected due to geometry. Diet or digestive physiology proxies did not have an influence on any intestinal length measures, though some proxies indicating more arid natural habitats were positively correlated with measures of the large intestine. The relative size of a forestomach compartment, the omasum, was negatively correlated with intestine length. The results indicate that intestine length measures provide little indication of feeding type or digestive physiology, but rather indicate adaptations to aridity. Higher-than-geometry scaling of intestinal length may be related to the necessity of maintaining geometric (or metabolic) scaling of intestinal surface area while keeping gut diameter, and hence the diffusion distances, small. The way in which space trade-offs determine the macroanatomy of different organs in the abdominal cavity, such as the omasum and the intestine, deserves further investigation.

Aspectos morfológicos do rúmen, omaso e fígado de tourinhos alimentados com diferentes níveis de glicerina bruta / Morphological aspects of rumen, omasum and liver of bulls fed different levels of crude glycerin

The morphology of the ruminal and omasal mucosa and hepatic parenchyma of 44 young bulls submitted to four types of concentrates were evaluated by varying the inclusion level of crude Glycerin - CG (0, 6, 12 and 18%). The variables evaluated were number of ruminal papillae per cm² of wall; area of the papillae and total absorption surface per cm² of wall; Mitotic index - MI, height and area of ruminal and omasal papillae; and hepatic morphology. The samples processed for inclusion in paraffin and stained with Hematoxylin-Eosin. The different levels of inclusion of CG in the diet did not interfere either in the number of papillae per cm2 of rumen wall (P= 0,70) or in the ruminal absorptive surface area (P= 0,89). Animals that received diets with levels 12 and 18% had higher MI of the ruminal epithelium (P< 0.01), but higher cell proliferation wasn't reflected in height gain and papillary area (P= 0,82). The omasal mucosa wasn't influenced by the inclusion of CG (MI e height gain and papillary area, P= 0,43, P= 0,56 e P= 0,9, respectively). The inclusion of GB in up to 18% of diet dry matter didn't alter the morphology of the rumen and omaso mucosa; and the integrity hepatic parenchyma.(AU)

Comparative omasum anatomy in ruminants: Relationships with natural diet, digestive physiology, and general considerations on allometric investigations.

The omasum is the third forestomach compartment of pecoran ruminants. It is assumed that the re-absorption of fluid present in the forestomach digesta (that facilitates particle sorting, digestion, and harvest of microbes) is its main function, so that less diluted digesta is submitted to enzymatic digestion in the lower digestive tract. Here, we evaluate measures of omasum size (representing 84 ruminant species in the largest data set) against body mass and proxies of the natural diet (%grass) or forestomach physiology (fluid throughput), using phylogenetically controlled models. The origin of specimens (free-ranging or captive) did not have an effect in the data set. Models with the best support invariably either included %grass or a physiology proxy in addition to body mass. These effects were not necessarily additive (affecting the intercept of the allometric regression), but often indicated a change in the allometric body mass-exponent with diet or physiology. Only models that allowed an influence on the allometric exponent yielded basic exponents compatible with predictions derived from geometry. Species that include more grass in their natural diet, or that have a "cattle-type" physiology marked by a high forestomach fluid throughput, generally have larger omasa. However, the existence of outliers, as well as the overall data pattern, suggest that this is not an obligatory morphophysiological condition. Circumstantial evidence is presented leading to the hypothesis that the comparatively small and less complex omasa of "moose-type" species do not necessarily represent an "original" state, but may be derived from more complex states by ontogenetic reduction and fusion of omasal laminae.

Molecular resolution to a morphological controversy: The case of North American fossil muskoxen Bootherium and Symbos.

The musk ox (Ovibos moschatus) is the only surviving member of a group of Pleistocene North American musk ox genera (Praeovibos, Ovibos, Bootherium, Euceratherium, and Soergelia) whose taxonomy is uncertain. The helmeted musk ox (Bootherium bombifrons) and the woodland musk ox (Symbos cavifrons) have been synonymised as male and female forms of a single Nearctic species found from Alaska, in the north, to Texas, in the south. However, this reclassification has not been tested using molecular data, despite the potential to use ancient DNA to examine these late Pleistocene taxa. In the present study, we sequenced mitochondrial genomes from seven subfossil musk ox specimens (originally identified as Bootherium and/or Symbos), allowing us to evaluate the identity of these muskoxen, explore their phylogeography, and estimate the timeline for their evolution. We also used nuclear genomic data to determine the sex of six of our seven samples. Ultimately, our molecular data support the synonymisation of the North American muskoxen Bootherium and Symbos.

Millions of Years Behind: Slow Adaptation of Ruminants to Grasslands.

The Late Cretaceous appearance of grasses, followed by the Cenozoic advancement of grasslands as dominant biomes, has contributed to the evolution of a range of specialized herbivores adapted to new diets, as well as to increasingly open and arid habitats. Many mammals including ruminants, the most diversified ungulate suborder, evolved high-crowned (hypsodont) teeth as an adaptation to tooth-wearing diets and habitats. The impact of different causes of tooth wear is still a matter of debate, and the temporal pattern of hypsodonty evolution in relation to the evolution of grasslands remains unclear. We present an improved time-calibrated molecular phylogeny of Cetartiodactyla, with phylogenetic reconstruction of ancestral ruminant diets and habitats, based on characteristics of extant taxa. Using this timeline, as well as the fossil record of grasslands, we conduct phylogenetic comparative analyses showing that hypsodonty in ruminants evolved as an adaptation to both diet and habitat. Our results demonstrate a slow, perhaps constrained, evolution of hypsodonty toward estimated optimal states, excluding the possibility of immediate adaptation. This augments recent findings that slow adaptation is not uncommon on million-year time scales.

Placental lesions associated with abortion and stillbirth in goats naturally infected by Neospora caninum / Lesões placentárias associadas a abortos e natimortos em cabras naturalmente infectadas pelo Neospora caninum

Neospora caninum has been described as a parasite that sporadically causes reproductive problems in goats. Several aspects of the pathogenesis of neosporosis in naturally infected goats remain to be established. The aims of the present study were to characterize the placental lesions in goats naturally infected by N. caninum and to evaluate several diagnostic techniques for effective detection of this protozoan in the goat placenta. Some placentas in this study originated from abortion and stillbirth in which there were severe lesions. The lesions were characterized mainly by necrosis involving the mesenchyme of the chorionic villi and trophoblast cells often alongside mononuclear inflammation and in some cases with neutrophilic infiltration. N. caninum DNA was detected in these placentas, but parasite structures were not visualized through immunohistochemistry (IHC). However, five of 11 placentas from N. caninum-infected goats that gave birth to healthy kids had histological lesions characterized by mononuclear inflammation. Of these 11 placentas, N. caninum DNA was detected in seven, and N. caninum tachyzoites were detected in only one of these seven placentas using IHC. The present study demonstrates that severe lesions in the placenta are associated with abortion and stillbirth in caprine neosporosis and the placental alterations are likely involved in abortion pathogenesis. Moreover, the results highlight the importance of using more than one diagnostic technique for the detection of the protozoan in placentas because N. caninum cannot be reliably detected by histological and immunohistochemical tests.(AU)

Neospora caninum é descrito como um parasito que causa problemas reprodutivos esporádicos em cabras. Muitos aspectos da patogênese da neosporose em cabras naturalmente infectadas ainda precisam ser estabelecidos. Os objetivos deste trabalho foram caracterizar as lesões placentárias em cabras naturalmente infectadas por N. caninum e avaliar as técnicas diagnósticas para a detecção efetiva do protozoário na placenta. Algumas placentas deste estudo são originárias de abortos e natimortos, nas quais havia lesões graves. As lesões foram classificadas principalmente por necrose envolvendo o mesênquima das vilosidades coriônicas e células trofoblásticas, geralmente associadas a infiltrado inflamatório mononuclear e em alguns casos infiltrado neutrofílico. O DNA do N. caninum foi detectado nestas placentas, porém estruturas parasitárias não foram visualizadas na imuno-histoquímica (IHQ). Entretanto, cinco das onze placentas de cabras infectadas, que deram à luz a conceptos saudáveis, apresentaram lesões histológicas caracterizadas por infiltrado inflamatório mononuclear. Destas 11 placentas, foi detectado DNA de N. caninum em sete e taquizoítos foram encontrados em apenas uma por meio de imuno-histoquímica. O presente estudo demonstra que abortos e natimortos na espécie caprina, causados pelo N. caninum estão associados a lesões acentuadas nas placentas, sendo que as mesmas estão envolvidas na patogênese do aborto. Os resultados também ressaltam a importância do uso de mais de uma técnica diagnóstica para a detecção do protozoário em placentas, pois o N. caninum não pode ser confiavelmente detectado somente pelos exames de histopatologia e imuno-histoquímica.(AU)

Micromeryx? eiselei-A new moschid species from Steinheim am Albuch, Germany, and the first comprehensive description of moschid cranial material from the Miocene of Central Europe.

Moschids are enigmatic pecoran ruminants whose phylogeny is still not fully understood. So far we know only little of the family's early evolutionary history and the origin of the modern genus, Moschus. Here we present a comprehensive description of cranial material, including the ear region and the dentition, of fossil moschid material from the Middle Miocene locality Steinheim am Albuch (13.5 Ma; Germany). This study provides the first exhaustive dataset for the cranial osteology of Micromeryx flourensianus, the most likely oldest true moschid. It furthermore reveals the presence of a second, so far undescribed moschid species, we here name Micromeryx? eiselei, in the abundant material from the locality. The two taxa can be clearly distinguished by characters of the skull, the ear region, the dentition, as well as by size. This evidences the sympatric occurrence of two moschid species in the locality Steinheim am Albuch.

Glycosylation and immunocytochemistry of binucleate cells in pronghorn (Antilocapra americana, Antilocapridae) show features of both Giraffidae and Bovidae.

Although the pronghorn (Antilocapra americana) resembles an antelope, its nearest relatives are the giraffe and okapi. In this study we have examined the placentae of 6 pronghorns using lectin- and immunocytochemistry to identify giraffid and bovid features. Binucleate cells (BNC) of the placenta exhibited features intermediate between those of the giraffe and bovine; Dolichos biflorus agglutinin binding - strong in the bovine BNC and absent in the giraffe - was evident in only a subpopulation of BNC while binding to blood vessels, as in the giraffe. Binding of Phytolacca americana agglutinin resembled that of the giraffe and okapi whereas many other glycans were found in all four clades. PAG antigens were similar to bovine and okapi but not giraffe. In summary, although the pronghorn outwardly resembles an antelope, placental BNC show giraffid features. Although each clade has its own individual characteristics, there are far more similarities than differences between them, emphasizing the common ancestry of all four clades.