The reduction in relative brain size in the domesticated dog is not an evolutionary singularity among the canids.

Domestication has long been considered the most powerful evolutionary engine behind dramatic reductions in brain size in several taxa, and the dog (Canis familiaris) is considered as a typical example that shows a substantial decrease in brain size relative to its ancestor, the grey wolf (Canis lupus). However, to make the case for exceptional evolution of reduced brain size under domestication requires an interspecific approach in a phylogenetic context that can quantify the extent by which domestication reduces brain size in comparison to closely related non-domesticated species responding to different selection factors in the wild. Here, we used a phylogenetic method to identify evolutionary singularities to test if the domesticated dog stands out in terms of relative brain size from other species of canids. We found that the dog does not present unambiguous signature of evolutionary singularity with regard to its small brain size, as the results were sensitive to the considerations about the ancestral trait values upon domestication. However, we obtained strong evidence for the hibernating common raccoon dog (Nyctereutes procyonoides) being an evolutionary outlier for its brain size. Therefore, domestication is not necessarily an exceptional case concerning evolutionary reductions in brain size in an interspecific perspective.

Rapid evolution of the primate larynx?

Tissue vibrations in the larynx produce most sounds that comprise vocal communication in mammals. Larynx morphology is thus predicted to be a key target for selection, particularly in species with highly developed vocal communication systems. Here, we present a novel database of digitally modeled scanned larynges from 55 different mammalian species, representing a wide range of body sizes in the primate and carnivoran orders. Using phylogenetic comparative methods, we demonstrate that the primate larynx has evolved more rapidly than the carnivoran larynx, resulting in a pattern of larger size and increased deviation from expected allometry with body size. These results imply fundamental differences between primates and carnivorans in the balance of selective forces that constrain larynx size and highlight an evolutionary flexibility in primates that may help explain why we have developed complex and diverse uses of the vocal organ for communication.

Selection and Constraints in the Ecomorphological Adaptive Evolution of the Skull of Living Canidae (Carnivora, Mammalia).

The association between phenotype and ecology is essential for understanding the environmental drivers of morphological evolution. This is a particularly challenging task when dealing with complex traits, such as the skull, where multiple selective pressures are at play and evolution might be constrained by ontogenetic and genetic factors. I integrate morphometric tools, comparative methods, and quantitative genetics to investigate how ontogenetic constraints and selection might have interacted during the evolution of the skull in extant Canidae. The results confirm that the evolution of cranial morphology was largely adaptive and molded by changes in diet composition. While the investigation of the adaptive landscape reveals two main selective lines of least resistance (one associated with size and one associated with functional shape features), rates of evolution along size were higher than those found for shape dimensions, suggesting the influence of constraints on morphological evolution. Structural modeling analyses revealed that size, which is the line of most genetic/phenotypic variation, might have acted as a constraint, negatively impacting dietary evolution. Constraints might have been overcome in the case of selection for the consumption of large prey by associating strong selection along both size and shape directions. The results obtained here show that microevolutionary constraints may have played a role in shaping macroevolutionary patterns of morphological evolution.

Ophthalmic contributions to assessing eyes of two neotropical canids: Cerdocyon thous and Chrysocyon brachyurus.

OBJECTIVE: To describe selected ophthalmic tests and anatomical features of eyes of crab-eating foxes (Cerdocyon thous) and maned wolves (Chrysocyon brachyurus). ANIMALS STUDIED: Six crab-eating foxes (12 eyes), eight maned wolves (16 eyes). PROCEDURES: Intramuscular and/or inhalatory anesthesia, ophthalmic evaluation with portable slit-lamp biomicroscope, Schirmer tear test (STT), intraocular pressure measurement with rebound and applanation (crab-eating fox only) tonometers, measurement of palpebral fissure length (PFL), gonioscopy, and fundoscopy. Data were analyzed with ANOVA and Tukey's tests. RESULTS: Both species presented upper and lower eyelids, both with eyelashes and Meibomian glands openings. A third eyelid was also present. In partial miosis, pigmented projections were observed along the edge of the pupil. The draining angle was open, with thin pectinate ligaments. The retina was holangiotic. For crab-eating foxes, mean ± standard deviation values were as follows: STT: 4.33 ± 2.96 mm/min; PFL: 17.45 ± 1.55 mm; rebound tonometry: 10.70 ± 3.43 mm Hg (TonoVet® calibration D),5.66 ± 3.44 mm Hg (TonoVet® calibration P), 17.00 ± 4.64 mm Hg (TonoVet® Plus calibration dog); and applanation tonometry: 11.70 ± 5.70 mm Hg(TonoPen® XL). For maned wolves, the mean ± standard deviation values were as follows: STT: 9.31 ± 7.40 mm/min; PFL: 22.79 ± 1.63 mm; rebound tonometry: 11.00 ± 2.77 mm Hg (TonoVet® calibration D), 6.78 ± 2.58 mm Hg (TonoVet® calibration P), and 18.29 ± 3.47 mm Hg(TonoVet® Plus calibration dog). CONCLUSIONS: This study contributes with knowledge that can help the clinical assessment regarding eyes of crab-eating foxes and maned wolves. The data herein presented for rebound tonometry are new for both species.

THE EYE OF CRAB-EATING FOX (CERDOCYON THOUS): ANATOMICAL CHARACTERISTICS AND NORMATIVE VALUES OF SELECTED DIAGNOSTIC TESTS, MORPHOMETRY OF CORNEAL TISSUE, AND ARRANGEMENTS OF CORNEAL STROMAL COLLAGEN FIBERS.

This study aimed to evaluate the ophthalmic parameters, morphometric features of corneal tissue, and arrangements of corneal stromal collagen fibers in crab-eating fox (Cerdocyon thous), a species of neotropical wild canid. We conducted the study on six juvenile crab-eating foxes (12 eyes), whilst 16 eyes were obtained post mortem from eight adult crab-eating foxes. The research was divided into two stages. In the first stage, eye anatomical characteristics, tear production (Schirmer 1 tear test, STT1), intraocular pressure (IOP), ocular echobiometry, and specular microscopy parameters related to morphology of corneal endothelium were studied in juvenile animals. In the second stage, morphometric features of corneal tissue (central corneal thickness [CCT] and corneal epithelium thickness) and arrangements of stromal collagen fibers were studied using eyes from adult animals. The main findings were that crab-eating fox eyes have vertical-slit pupils, holangiotic retina, and reference values (mean ± SD) of 13.37 ± 3.79 mm/min for STT1 and of 10.43 ± 3.84 mmHg for IOP. The ocular echobiometric features observed in crab-eating foxes are different from those reported for domestic dogs (Canis familiaris). Conversely, the corneal endothelial parameters are similar to those of domestic dogs. The CCT measured by tissue morphometry was 0.54 ± 0.06 mm, and the corneal epithelium thickness was 60.13 ± 8.71 µm. Mean coherency related to alignment of collagen fibers was 0.66 ± 0.12. The crab-eating fox cornea had predominantly thick collagen fibers. Crab-eating fox eyes have morphofunctional peculiarities. They resemble the eyes of domestic dogs in some aspects, but diverge in others.

The late Neopleistocene dhole (Carnivora, Canidae, Cuon alpinus Pallas, 1811) from the Urals.

Morphotypic and morphometric analyses of the M2 tooth found in the Late Pleistocene deposits of the Ignatievskaya Cave (southern Urals) demonstrate that this tooth belongs to the dhole (Cuon alpinus Pallas, 1811). This is the first reliable evidence for the presence of the dhole in the Urals. Radiocarbon dating and associated theriofauna allow the tooth to be dated to the first half of the late Neopleistocene or, more precisely, to marine isotope stages (MISs) 3 and 4.

Brainy stuff of long-gone dogs: a reappraisal of the supposed Canis endocranial cast from the Pliocene of Poland.

The pre-Quaternary fossil record of Canis in the Old World is scarce, and the first appearance of this genus in Europe remains an enigma. Amongst the oldest fossils assigned to this genus, there is a natural cast of the brain (endocast) collected in Weze 1, Poland, from Pliocene deposits dated between 3.3 and 4.0 Ma. We reexamined this specimen and found that it differs from the brain of Canis in having its region medial to the coronal sulcus heart-shaped in dorsal view, its region rostral to the presylvian sulcus shorter and less constricted laterally, and its cerebellum less overlapped by the cerebrum and lacking a lateral twist of the posterior vermis. We identified this fossil, as well as another fossil canid endocast from Weze 1, as representing the raccoon dog genus Nyctereutes. The previously reported presence of Canis in Weze 1 is therefore not confirmed. Specifically, both endocasts can be referred to N. donnezani because this is the only species of Nyctereutes that has been recognised in this locality on the basis of craniomandibular and dental fossils. Our study represents a taxonomic application of comparative neuroanatomical and palaeoneurological data, an approach that may become increasingly useful with the growing knowledge of the endocranial morphology of fossil mammals.

High-resolution computed tomographic (HRCT) image series from 413 canid and 18 felid skulls.

Computed tomography (CT) is a non-invasive, three-dimensional imaging tool used in medical imaging, forensic science, industry and engineering, anthropology, and archaeology. The current study used high-resolution medical CT scanning of 431 animal skulls, including 399 dog skulls from 152 breeds, 14 cat skulls from 9 breeds, 14 skulls from 8 wild canid species (gray wolf, golden jackal, coyote, maned wolf, bush dog, red fox, Fennec fox, bat-eared fox), and 4 skulls from 4 wild felid species (wildcat, leopard, serval, caracal). This comprehensive and unique collection of CT image series of skulls can provide a solid foundation not only for comparative anatomical and evolutionary studies but also for the advancement of veterinary education, virtual surgery planning, and the facilitation of training in sophisticated machine learning methodologies.

Comparative anatomy of the caudate nucleus in canids and felids: Associations with brain size, curvature, cross-sectional properties, and behavioral ecology.

The evolutionary history of canids and felids is marked by a deep time separation that has uniquely shaped their behavior and phenotype toward refined predatory abilities. The caudate nucleus is a subcortical brain structure associated with both motor control and cognitive, emotional, and executive functions. We used a combination of three-dimensional imaging, allometric scaling, and structural analyses to compare the size and shape characteristics of the caudate nucleus. The sample consisted of MRI scan data obtained from six canid species (Canis lupus lupus, Canis latrans, Chrysocyon brachyurus, Lycaon pictus, Vulpes vulpes, Vulpes zerda), two canid subspecies (Canis lupus familiaris, Canis lupus dingo), as well as three felids (Panthera tigris, Panthera uncia, Felis silvestris catus). Results revealed marked conservation in the scaling and shape attributes of the caudate nucleus across species, with only slight deviations. We hypothesize that observed differences in caudate nucleus size and structure for the domestic canids are reflective of enhanced cognitive and emotional pathways that possibly emerged during domestication.

Radiographic evaluation of the forelimb bone development in maned wolves (Chrysocyon brachyurus).

This study aimed to assess the fusion of growth plates and the development of secondary ossification centres in the forelimb bones of maned wolves (Chrysocyon brachyurus), contrasting the findings with established data from domestic dogs. Three maned wolves, comprising one male and two females, initially aged between 3 and 4 months, were subjected to monthly radiographic evaluations until 10-11 months of age, followed by bimonthly assessments until 18-19 months of age, encompassing both forelimbs. The closure times of growth plates were observed as follows: supraglenoid tubercle (7-8 months), proximal humerus (17-19 months), distal humerus (8-9 months), medial epicondyle of the humerus (8-9 months), proximal ulna (9-10 months), proximal radius (13-15 months), distal ulna (13-15 months) and distal radius (17-19 months). Statistical analysis revealed significant differences in the areas of secondary ossification centres in the proximal epiphyses of the humerus and radius, respectively, observed from the initial evaluation at 8-9 months and 6-7 months. Conversely, the epiphyses of the supraglenoid tubercle, distal humerus, proximal ulna, distal ulna, medial epicondyle of the humerus and distal radius did not exhibit significant area differences between 3-4 months and 4-5 months, yet notable distinctions emerged at 5-6 months. In summary, while the radiographic appearance of epiphyseal growth plates and secondary ossification centres in maned wolves resembles that of domestic dogs, closure times vary. These findings contribute to understanding the dynamics of epiphyseal growth plates in this species.

A pathological Late Pleistocene canid from San Sidero (Italy): implications for social- and feeding-behaviour.

Evidence of diseases on vertebrate fossil bones can provide detailed information on many aspects of extinct animals. This study focused on pathological craniodental remains (left maxilla and dentary) referred to the canid Cuon alpinus unearthed from a Late Pleistocene karst filling deposit at San Sidero (Apulia, southern Italy). These fossils show clear evidence of a chronic periodontitis that caused the animal's death. Clinical diagnosis of the disease and the timing of its development have been defined on the basis of a veterinary odontostomatology approach, in addition to radiographic and tomographic techniques. From the initiation of the infection until death, a time span of at least 6 months occurred, and three main steps have been defined: (1) the bacterial infections of the buccal cavity turning into severe periodontitis, (2) the fracture of the lower carnassial and (3) the loss of teeth due to the worsening infection that deformed and/or eroded maxillary and mandibular bones and enlarged alveoli. The analysis of the palaeopathology also provides information about the biomechanics of the bite, on the feeding behaviour and on the relationships of injured members in a pack of Late Pleistocene canids.

Comparative analysis of origin and branches of phrenic nerve in the diaphragm of the crab-eating fox (Cerdocyon thous) and domestic dog (Canis lupus familiaris).

The crab-eating fox (Cerdocyon thous) is a canid widely distributed throughout South America, considered the only representative of the gender. There is a lack of information about anatomical aspects of the species, mainly regarded of nervous system anatomy. For clinical and surgical approaches of these animals, the domestic dog is adopted as anatomophysiological model. So, the aim of the present study was to analyse origin and branches of the phrenic nerve in the diaphragm of crab-eating fox compared to domestic dog. Four specimens of Cerdocyon thous and four of Canis lupus familiaris were used. The phrenic nerve originated from the ventral branches of the fifth (C5), sixth (C6) and seventh (C7) cervical spinal nerves. Phrenic nerves were distributed in the diaphragm as lumbocostal trunk and sternal branch (100%) in the crab-eating fox, and in costosternal trunk and lumbar branch (75%) and lumbocostal trunk and sternal branch (25%) in the domestic dog. In both species, the lumbar branch innervates the diaphragm pillars, the costal branch addresses all costal region, and the sternal branch distributes in the costal ventrolateral region and the sternal part of the diaphragm, to the left and to the right.

Dispelling dog dogma: an investigation of heterochrony in dogs using 3D geometric morphometric analysis of skull shape.

Heterochrony is an evolutionary mechanism that generates diversity via perturbations of the rate or timing of development that requires very little genetic innovation. As such, heterochrony is thought to be a common evolutionary mechanism in the generation of diversity. Previous research has suggested that dogs evolved via heterochrony and are paedomorphic wolves. This study uses three-dimensional landmark-based coordinate data to investigate heterochronic patterns within the skull morphology of the domestic dog. A total of 677 adult dogs representing 106 different breeds were measured and compared with an ontogenetic series of 401 wolves. Geometric morphometric analysis reveals that the cranial shape of none of the modern breeds of dogs resembles the cranial shapes of adult or juvenile wolves. In addition, investigations of regional heterochrony in the face and neurocranium also reject the hypothesis of heterochrony. Throughout wolf cranial development the position of the face and the neurocranium remain in the same plane. Dogs, however, have a de novo cranial flexion in which the palate is tilted dorsally in brachycephalic and mesaticephalic breeds or tilted ventrally in dolichocephalic and down-face breeds. Dogs have evolved very rapidly into an incredibly morphologically diverse species with very little genetic variation. However, the genetic alterations to dog cranial development that have produced this vast range of phylogenetically novel skull shapes do not coincide with the expectations of the heterochronic model. Dogs are not paedomorphic wolves.

Volumetric assessment of cerebral asymmetries in dogs.

In the present study we quantified volumetric brain asymmetries from computed tomography (CT) scans in 12 healthy dogs, using a semi-automated technique for assessing in vivo structure asymmetry. Volumetric assessment of asymmetrical cerebral lateral ventricle (ALV) was also investigated. Our results showed that seven dogs exhibited a right hemisphere significantly greater than the left, two dogs had a left-greater-than-right hemisphere asymmetry, and finally two dogs displayed no significant brain volumetric asymmetry. This right-biased hemispheric asymmetry supports data reported previously using post-mortem morphological studies in both dogs and other mammalian species.

Anatomical variations in cervical vertebrae in two species of neotropical canids: What is the meaning?

Anatomical variation in wild canids is rarely described. Altered shapes, uni- or bilaterally situated, were observed in ventral laminae of C6 and/or in C7 vertebral body of two fox species: Lycalopex gymnocercus and Cerdocyon thous. The specimens were young adults, collected dead on highways in southern Brazil. Deformities were considered anatomical variations because apparently there was no interference in function. The occurrence of such variations in two related species from the same area can suggest an environmental common cause rather than genetic issues. Future studies may deepen the understanding between cause and effect of these vertebral deformities in wild canids.

Anatomical and morphometric evaluation of the orbit, eye tunics, eyelids and orbital glands of the captive females of the South African painted dog (Lycaon pictus pictus Temminck, 1820) (Caniformia: Canidae).

In this study, we present the first data concerning the anatomical, morphometrical, histological and histochemical study of the orbit, eye tunics, eyelids and orbital glands in South African Painted Dogs (Lycaon pictus pictus). The study was performed using eyeball morphometry, analysis of the bony orbit including its morphometry, macroscopic study, morphometry, histological examination of the eye tunics and chosen accessory organs of the eye and histochemical analysis. The orbit was funnel shaped and was open-type. There was a single ethmoid opening for the ethmoid nerve on the orbital lamina. The pupil was round, while the ciliary body occupied a relatively wide zone. The iris was brown and retina had a pigmented area. The cellular tapetum lucidum was semi-circular and milky and was composed of 14-17 layers of tapetal cells arranged in a bricklike structure. In the lower eyelid, there was a single conjunctival lymph nodule aggregate. One or two additional large conjunctval folds were observed within the posterior surface of the upper eyelids. The superficial gland of the third eyelid had a serous nature. The third eyelid was T-shaped and was composed of hyaline tissue. Two to three conjunctival lymph nodul aggregates were present within the bulbar conjunctiva of the third eyelid. The lacrimal gland produced a sero-mucous secretion. A detailed anatomic analysis of the eye area in the captive South African Painted Dogs females showed the similarities (especially in the histological examination of the eyetunics and orbital glands) as well as the differences between the Painted dog and the other representatives of Canidae. The differences included the shape and size od the orbita with comparison to the domestic dog. Such differences in the orbit measurements are most likely associated with the skull type, which are defined in relation to domestic dogs. The presented results significantly expand the existing knowledge on comparative anatomy in the orbit, eye and chosen accessory organs in wild Canidae.

Ancient DNA from the Asiatic Wild Dog (Cuon alpinus) from Europe.

The Asiatic wild dog (Cuon alpinus), restricted today largely to South and Southeast Asia, was widespread throughout Eurasia and even reached North America during the Pleistocene. Like many other species, it suffered from a huge range loss towards the end of the Pleistocene and went extinct in most of its former distribution. The fossil record of the dhole is scattered and the identification of fossils can be complicated by an overlap in size and a high morphological similarity between dholes and other canid species. We generated almost complete mitochondrial genomes for six putative dhole fossils from Europe. By using three lines of evidence, i.e., the number of reads mapping to various canid mitochondrial genomes, the evaluation and quantification of the mapping evenness along the reference genomes and phylogenetic analysis, we were able to identify two out of six samples as dhole, whereas four samples represent wolf fossils. This highlights the contribution genetic data can make when trying to identify the species affiliation of fossil specimens. The ancient dhole sequences are highly divergent when compared to modern dhole sequences, but the scarcity of dhole data for comparison impedes a more extensive analysis.

The early hunting dog from Dmanisi with comments on the social behaviour in Canidae and hominins.

The renowned site of Dmanisi in Georgia, southern Caucasus (ca. 1.8 Ma) yielded the earliest direct evidence of hominin presence out of Africa. In this paper, we report on the first record of a large-sized canid from this site, namely dentognathic remains, referable to a young adult individual that displays hypercarnivorous features (e.g., the reduction of the m1 metaconid and entoconid) that allow us to include these specimens in the hypodigm of the late Early Pleistocene species Canis (Xenocyon) lycaonoides. Much fossil evidence suggests that this species was a cooperative pack-hunter that, unlike other large-sized canids, was capable of social care toward kin and non-kin members of its group. This rather derived hypercarnivorous canid, which has an East Asian origin, shows one of its earliest records at Dmanisi in the Caucasus, at the gates of Europe. Interestingly, its dispersal from Asia to Europe and Africa followed a parallel route to that of hominins, but in the opposite direction. Hominins and hunting dogs, both recorded in Dmanisi at the beginning of their dispersal across the Old World, are the only two Early Pleistocene mammal species with proved altruistic behaviour towards their group members, an issue discussed over more than one century in evolutionary biology.

Little brainiacs and big dummies: Are we selecting for stupid, stout, or small dogs?

Background: Brain size has been associated with intelligence of various orders and families of animals, leading to the concept of encephalization. Brain size scales with body weight between species within mammals to approximately the 0.67 power. However, within species, this scaling exponent appears to be much smaller (approximately 0.27 power). Aim: We examined whether this relationship has persisted in dogs over the 120 years since this was originally observed. Methods: Comparative cross-sectional study of magnetic resonance imaging (MRI) data obtained from 127 dogs, compared to historical data from 157 dogs and 24 non-dog canid species. Results: Brain size in dogs measured by MRI had a scaling exponent virtually identical to that observed previously (0.24 vs. 0.26). However, the proportionality constant was smaller, suggesting that dogs in the study cohort had relatively smaller brains than the historical cohort. Absolute brain size appeared to have both a lower and upper limit in dogs. When compared to non-dogs canids, the most appropriate "representative" size for a "typical dog" when examining allometric scaling across Canidae appeared to be approximately 10-15 kg. Conclusions: We interpreted the slight reduction in relative brain size to be a function of increased obesity in the study cohort compared to dogs examined 120 years ago. Further, we suggest that dog brains have a finite lower size limit. Finally, concepts of encephalization should not be applied to dogs.

The evolution of South American endemic canids: a history of rapid diversification and morphological parallelism.

The origin of endemic South American canid fauna has been traditionally linked with the rise of the Isthmus of Panama, suggesting that diversification of the dog fauna on this continent occurred very rapidly. Nevertheless, despite its obvious biogeographic appeal, the tempo of Canid evolution in South America has never been studied thoroughly. This issue can be suitably tackled with the inference of a molecular timescale. In this study, using a relaxed molecular clock method, we estimated that the most recent common ancestor of South American canids lived around 4 Ma, whereas all other splits within the clade occurred after the rise of the Panamanian land bridge. We suggest that the early diversification of the ancestors of the two main lineages of South American canids may have occurred in North America, before the Great American Interchange. Moreover, a concatenated morphological and molecular analysis put some extinct canid species well within the South American radiation, and shows that the dental adaptations to hypercarnivory evolved only once in the South American clade.