Teeth out of proportion: Smaller horse and cattle breeds have comparatively larger teeth.

There are different descriptions of allometric relationships between important components of the mammalian skull. Craniofacial evolutionary allometry describes a pattern of increasing facial cranium in larger skulls. Another body of literature describes disproportionately larger teeth in smaller species or specimens, matching anecdotal observations with dental problems in dwarf breeds whose teeth appear "too large for their skulls." We test the scaling of tooth row length with body size and skull length in a data set comprising 114 domestic horses (representing 40 breeds) and in another data set of 316 domestic cattle (of >60 breeds). We demonstrate that smaller skulls have a relatively longer tooth row in both horses and cattle; larger specimens have relatively shorter tooth rows. Whereas in horses, larger skulls have a relatively longer diastema, the distance of the mesial maxillary premolar to the premaxilla was proportional to cranium length in cattle. While the reasons for these patterns remain to be detected, they support the hypothesis that tooth size might be less "evolvable," in terms of time required for changes, than body size. The pattern may affect (i) the selective breeding for dwarf breeds by setting minimum constraints for skull size, as described previously for domestic horses with the same data set; (ii) the susceptibility of small breeds for dental problems; and (iii) differences in chewing efficiency between breeds of different sizes. The findings support the existing concept that scaling of tooth to body size across taxa becomes more isometric the longer these taxa are separated in evolutionary time.

Cranial shape variation in domestication: A pilot study on the case of rabbits.

Domestication leads to phenotypic characteristics that have been described to be similar across species. However, this "domestication syndrome" has been subject to debate, related to a lack of evidence for certain characteristics in many species. Here we review diverse literature and provide new data on cranial shape changes due to domestication in the European rabbit (Oryctolagus cuniculus) as a preliminary case study, thus contributing novel evidence to the debate. We quantified cranial shape of 30 wild and domestic rabbits using micro-computed tomography scans and three-dimensional geometric morphometrics. The goal was to test (1) if the domesticates exhibit shorter and broader snouts, smaller teeth, and smaller braincases than their wild counterparts; (2) to what extent allometric scaling is responsible for cranial shape variation; (3) if there is evidence for more variation in the neural crest-derived parts of the cranium compared with those derived of the mesoderm, in accordance with the "neural crest hypothesis." Our own data are consistent with older literature records, suggesting that although there is evidence for some cranial characteristics of the "domestication syndrome" in rabbits, facial length is not reduced. In accordance with the "neural crest hypothesis," we found more shape variation in neural crest versus mesoderm-derived parts of the cranium. Within the domestic group, allometric scaling relationships of the snout, the braincase, and the teeth shed new light on ubiquitous patterns among related taxa. This study-albeit preliminary due to the limited sample size-adds to the growing evidence concerning nonuniform patterns associated with domestication.

Similar rates of morphological evolution in domesticated and wild pigs and dogs.

BACKGROUND: Whether the great morphological disparity of domesticated forms is the result of uniformly higher evolutionary rates compared to the wild populations is debated. We provide new data on changes of skull dimensions within historical time periods in wild and domesticated dogs and pigs to test if domestication might lead to an accelerated tempo of evolution in comparison to the wild conspecifics. Darwins and Haldanes were used to quantify evolutionary rates. Comparisons with evolutionary rates in other species and concerning other characteristics from the literature were conducted. RESULTS: Newly gathered and literature data show that most skull dimensions do not change faster in domesticated breeds than in wild populations, although it is well known that there is extensive artificial selection on skull shape in some dog breeds. Evolutionary rates among domesticated forms and traits (e.g., production traits in pigs, and racing speed in some horses and greyhounds) might vary greatly with species and breeding aim. CONCLUSIONS: Our study shows that evolutionary rates in domestication are not in any event faster than those in the wild, although they are often perceived as such given the vast changes that appear in a relatively short period of time. This may imply that evolution under natural conditions - i.e., without human intervention - is not as slow as previously described, for example by Darwin. On the other hand, our results illustrate how diverse domestication is in tempo, mode, and processes involved.

Geographic variation in responses of European yellow dung flies to thermal stress.

Climatic conditions can be very heterogeneous even over small geographic scales, and are believed to be major determinants of the abundance and distribution of species and populations. Organisms are expected to evolve in response to the frequency and magnitude of local thermal extremes, resulting in local adaptation. Using replicate yellow dung fly (Scathophaga stercoraria; Diptera: Scathophagidae) populations from cold (northern Europe) and warm climates (southern Europe), we compared 1) responses to short-term heat and cold shocks in both sexes, 2) heat shock protein (Hsp70) expression in adults and eggs, and 3) female reproductive traits when facing short-term heat stress during egg maturation. Contrary to expectations, thermal traits showed minor geographic differentiation, with weak evidence for greater heat resistance of southern flies but no differentiation in cold resistance. Hsp70 protein expression was little affected by heat stress, indicating systemic rather than induced regulation of the heat stress response, possibly related to this fly group's preference for cold climes. In contrast, sex differences were pronounced: males (which are larger) endured hot temperatures longer, while females featured higher Hsp70 expression. Heat stress negatively affected various female reproductive traits, reducing first clutch size, overall reproductive investment, egg lipid content, and subsequent larval hatching. These responses varied little across latitude but somewhat among populations in terms of egg size, protein content, and larval hatching success. Several reproductive parameters, but not Hsp70 expression, exhibited heritable variation among full-sib families. Rather than large-scale clinal geographic variation, our study suggests some local geographic population differentiation in the ability of yellow dung flies to buffer the impact of heat stress on reproductive performance.

Growth trajectories in the cave bear and its extant relatives: an examination of ontogenetic patterns in phylogeny.

BACKGROUND: The study of postnatal ontogeny can provide insights into evolution by offering an understanding of how growth trajectories have evolved resulting in adult morphological disparity. The Ursus lineage is a good subject for studying cranial and mandibular shape and size variation in relation to postnatal ontogeny and phylogeny because it is at the same time not diverse but the species exhibit different feeding ecologies. Cranial and mandibular shapes of Ursus arctos (brown bear), U. maritimus (polar bear), U. americanus (American black bear), and the extinct U. spelaeus (cave bear) were examined, using a three-dimensional geometric morphometric approach. Additionally, ontogenetic series of crania and mandibles of U. arctos and U. spelaeus ranging from newborns to senile age were sampled. RESULTS: The distribution of specimens in morphospace allowed to distinguish species and age classes and the ontogenetic trajectories U. arctos and U. spelaeus were found to be more similar than expected by chance. Cranial shape changes during ontogeny are largely size related whereas the evolution of cranial shape disparity in this clade appears to be more influenced by dietary adaptation than by size and phylogeny. The different feeding ecologies are reflected in different cranial and mandibular shapes among species. CONCLUSIONS: The cranial and mandibular shape disparity in the Ursus lineage appears to be more influenced by adaptation to diet than by size or phylogeny. In contrast, the cranial and mandibular shape changes during postnatal ontogeny in U. arctos and U. spelaeus are probably largely size related. The patterns of morphospace occupation of the cranium and the mandible in adults and through ontogeny are different.

Correction: 'A longitudinal study of phenotypic changes in early domestication of house mice' (2018), by Geiger et al.

[This corrects the article DOI: 10.1098/rsos.172099.][This corrects the article DOI: 10.1098/rsos.172099.].

Skeletal variation in bird domestication: limb proportions and sternum in chicken, with comparisons to mallard ducks and Muscovy ducks.

Background: Domestication, including selective breeding, can lead to morphological changes of biomechanical relevance. In birds, limb proportions and sternum characteristics are of great importance and have been studied in the past for their relation with flight, terrestrial locomotion and animal welfare. In this work we studied the effects of domestication and breed formation in limb proportions and sternum characteristics in chicken (Gallus gallus), mallard ducks (Anas plathyrhynchos) and Muscovy ducks (Cairina moschata). Methods: First, we quantified the proportional length of three long bones of the forelimb (humerus, radius, and carpometacarpus) and the hind limb (femur, tibiotarsus, and tarsometatarsus) in domestic chickens, mallard ducks, and Muscovy ducks and their wild counterparts. For this, we took linear measurements of these bones and compared their proportions in the wild vs. the domestic group in each species. In chicken, these comparisons could also be conducted among different breeds. We then evaluated the proportional differences in the context of static and ontogenetic allometry. Further, we compared discrete sternum characteristics in red jungle fowl and chicken breeds. In total, we examined limb bones of 287 specimens and keel bones of 63 specimens. Results: We found a lack of significant change in the proportions of limb bones of chicken and Muscovy duck due to domestication, but significant differences in the case of mallard ducks. Variation of evolvability, allometric scaling, and heterochrony may serve to describe some of the patterns of change we report. Flight capacity loss in mallard ducks resulting from domestication may have a relation with the difference in limb proportions. The lack of variation in proportions that could distinguish domestic from wild forms of chicken and Muscovy ducks may reflect no selection for flight capacity during the domestication process in these groups. In chicken, some of the differences identified in the traits discussed are breed-dependent. The study of the sternum revealed that the condition of crooked keel was not unique to domestic chicken, that some sternal characteristics were more frequent in certain chicken breeds than in others, and that overall there were no keel characteristics that are unique for certain chicken breeds. Despite some similar morphological changes identified across species, this study highlights the lack of universal patterns in domestication and breed formation.

Pattern and pace of morphological change due to variable human impact: the case of Japanese macaques.

Human impact influences morphological variation in animals, as documented in many captive and domestic animal populations. However, there are different levels of human impact, and their influence on the pattern and rate of morphological variation remains unclear. This study contributes to the ongoing debate via the examination of cranial and mandibular shape and size variation and pace of change in Japanese macaques (Macaca fuscata). This species is ideal for tackling such questions because different wild, wild-provisioned, and captive populations have been monitored and collected over seven decades. Linear measurements were taken on 70 skulls from five populations, grouped into three 'human impact groups' (wild, wild-provisioned, and captive). This made it possible to investigate the pattern and pace of skull form changes among the human impact groups as well as over time within the populations. It was found that the overall skull shape tends to differ among the human impact groups, with captive macaques having relatively longer rostra than wild ones. Whether these differences are a result of geographic variation or variable human impact, related to nutritional supply and mechanical properties of the diet, is unclear. However, this pattern of directed changes did not seem to hold when the single captive populations were examined in detail. Although environmental conditions have probably been similar for the two examined captive populations (same captive locality), skull shape changes over the first generations in captivity were mostly different. This varying pattern, together with a consistent decrease in body size in the captive populations over generations, points to genetic drift playing a role in shaping skull shape and body size in captivity. In the captive groups investigated here, the rates of change were found to be high compared to literature records from settings featuring different degrees of human impact in different species, although they still lie in the range of field studies in a natural context. This adds to the view that human impact might not necessarily lead to particularly fast rates of change.

Modularity patterns in mammalian domestication: Assessing developmental hypotheses for diversification.

The neural crest hypothesis posits that selection for tameness resulted in mild alterations to neural crest cells during embryonic development, which directly or indirectly caused the appearance of traits associated with the "domestication syndrome" (DS). Although representing an appealing unitary explanation for the generation of domestic phenotypes, support for this hypothesis from morphological data and for the validity of the DS remains a topic of debate. This study used the frameworks of morphological integration and modularity to assess patterns that concern the embryonic origin of the skull and issues around the neural crest hypothesis. Geometric morphometric landmarks were used to quantify cranial trait interactions between six pairs of wild and domestic mammals, comprising representatives that express between five and 17 of the traits included in the DS, and examples from each of the pathways by which animals entered into relationships with humans. We predicted the presence of neural crest vs mesoderm modular structure to the cranium, and that elements in the neural crest module would show lower magnitudes of integration and higher disparity in domestic forms compared to wild forms. Our findings support modular structuring based on tissue origin (neural crest, mesoderm) modules, along with low module integration magnitudes for neural crest cell derived cranial elements, suggesting differential capacity for evolutionary response among those elements. Covariation between the neural crest and mesoderm modules accounted for major components of shape variation for most domestic/wild pairs. Contra to our predictions, however, we find domesticates share similar integration magnitudes to their wild progenitors, indicating that higher disparity in domesticates is not associated with magnitude changes to integration among either neural crest or mesoderm derived elements. Differences in integration magnitude among neural crest and mesoderm elements across species suggest that developmental evolution preserves a framework that promotes flexibility under the selection regimes of domestication.

Decline in Distribution and Abundance: Urban Hedgehogs under Pressure.

Increasing urbanization and densification are two of the largest global threats to biodiversity. However, certain species thrive in urban spaces. Hedgehogs Erinaceus europaeus have been found in higher densities in green areas of settlements as compared to rural spaces. With recent studies pointing to dramatically declining hedgehog numbers in rural areas, we pose the question: how do hedgehogs fare in urban spaces, and do these spaces act as refuges? In this study, recent (2016-2018) and past (1992) hedgehog abundance and distribution were compared across the city of Zurich, Switzerland using citizen science methods, including: footprint tunnels, capture-mark recapture, and incidental sightings. Our analyses revealed consistent negative trends: Overall hedgehog distribution decreased by 17.6% ± 4.7%, whereas abundance declined by 40.6% (mean abundance 32 vs. 19 hedgehogs/km2, in past and recent time, respectively), with one study plot even showing a 91% decline in this period (78 vs. 7 hedgehogs/km2, respectively). We discuss possible causes of this rapid decline: increased urban densification, reduction of insect biomass, and pesticide use, as well as the role of increasing populations of badgers (a hedgehog predator) and parasites or diseases. Our results suggest that hedgehogs are now under increasing pressure not only in rural but also in urban areas, their former refuges.

A longitudinal study of phenotypic changes in early domestication of house mice.

Similar phenotypic changes occur across many species as a result of domestication, e.g. in pigmentation and snout size. Experimental studies of domestication have concentrated on intense and directed selection regimes, while conditions that approximate the commensal and indirect interactions with humans have not been explored. We examine long-term data on a free-living population of wild house mice that have been indirectly selected for tameness by regular exposure to humans. In the course of a decade, this mouse population exhibited significantly increased occurrence of white patches of fur and decreased head length. These phenotypic changes fit to the predictions of the 'domestication syndrome'.

Neomorphosis and heterochrony of skull shape in dog domestication.

The overall similarity of the skull shape of some dog breeds with that of juvenile wolves begs the question if and how ontogenetic changes such as paedomorphosis (evolutionary juvenilisation) played a role in domestication. Here we test for changes in patterns of development and growth during dog domestication. We present the first geometric morphometric study using ontogenetic series of dog and wolf crania, and samples of dogs with relatively ancestral morphology and from different time periods. We show that patterns of juvenile-to-adult morphological change are largely similar in wolves and domestic dogs, but differ in two ways. First, dog skulls show unique (neomorphic) features already shortly after birth, and these features persist throughout postnatal ontogeny. Second, at any given age, juvenile dogs exhibit skull shapes that resemble those of consistently younger wolves, even in dog breeds that do not exhibit a 'juvenilized' morphology as adults. These patterns exemplify the complex nature of evolutionary changes during dog domestication: the cranial morphology of adult dogs cannot simply be explained as either neomorphic or paedomorphic. The key to our understanding of dog domestication may lie in a closer comparative examination of developmental phases.

On the lack of a universal pattern associated with mammalian domestication: differences in skull growth trajectories across phylogeny.

As shown in a taxonomically broad study, domestication modifies postnatal growth. Skull shape across 1128 individuals was characterized by 14 linear measurements, comparing 13 pairs of wild versus domesticated forms. Among wild forms, the boar, the rabbit and the wolf have the highest proportion of allometric growth, explaining in part the great morphological diversity of the domesticated forms of these species. Wild forms exhibit more isometric growth than their domesticated counterparts. Multivariate comparisons show that dogs and llamas exhibit the greatest amount of differences in trajectories with their wild counterparts. The least amount is recorded in the pig-boar, and camel and horse pairs. Bivariate analyses reveal that most domesticated forms have growth trajectories different from their respective wild counterparts with regard to the slopes. In pigs and camels slopes are shared and intercepts are different. There is a trajectory extension in most domesticated herbivores and the contrary pattern in carnivorous forms. However, there is no single, universal and global pattern of paedomorphosis or any other kind of heterochrony behind the morphological diversification that accompanies domestication.

Cranial Suture Closure in Domestic Dog Breeds and Its Relationships to Skull Morphology.

Bulldog-type brachycephalic domestic dog breeds are characterized by a relatively short and broad skull with a dorsally rotated rostrum (airorhynchy). Not much is known about the association between a bulldog-type skull conformation and peculiar patterns of suture and synchondrosis closure in domestic dogs. In this study, we aim to explore breed-specific patterns of cranial suture and synchondrosis closure in relation to the prebasial angle (proxy for airorhynchy and thus bulldog-type skull conformation) in domestic dogs. For this purpose, we coded closure of 18 sutures and synchondroses in 26 wolves, that is, the wild ancestor of all domestic dogs, and 134 domestic dogs comprising 11 breeds. Comparisons of the relative amount of closing and closed sutures and synchondroses (closure scores) in adult individuals showed that bulldog-type breeds have significantly higher closure scores than non-bulldog-type breeds and that domestic dogs have significantly higher closure scores than the wolf. We further found that the prebasial angle is significantly positively correlated with the amount of closure of the basispheno-presphenoid synchondrosis and sutures of the nose (premaxillo-nasal and maxillo-nasal) and the palate (premaxillo-maxillary and interpalatine). Our results show that there is a correlation between patterns of suture and synchondrosis closure and skull shape in domestic dogs, although the causal relationships remain elusive.

The taming of the neural crest: a developmental perspective on the origins of morphological covariation in domesticated mammals.

Studies on domestication are blooming, but the developmental bases for the generation of domestication traits and breed diversity remain largely unexplored. Some phenotypic patterns of human neurocristopathies are suggestive of those reported for domesticated mammals and disrupting neural crest developmental programmes have been argued to be the source of traits deemed the 'domestication syndrome'. These character changes span multiple organ systems and morphological structures. But an in-depth examination within the phylogenetic framework of mammals including domesticated forms reveals that the distribution of such traits is not universal, with canids being the only group showing a large set of predicted features. Modularity of traits tied to phylogeny characterizes domesticated mammals: through selective breeding, individual behavioural and morphological traits can be reordered, truncated, augmented or deleted. Similarly, mammalian evolution on islands has resulted in suites of phenotypic changes like those of some domesticated forms. Many domesticated mammals can serve as valuable models for conducting comparative studies on the evolutionary developmental biology of the neural crest, given that series of their embryos are readily available and that their phylogenetic histories and genomes are well characterized.

Unaltered sequence of dental, skeletal, and sexual maturity in domestic dogs compared to the wolf.

BACKGROUND: It has been hypothesised that domestication altered the sequence of dental, skeletal, and sexual maturity of dogs when compared to their wolf ancestor. To test this we investigated a comprehensive sample of domestic dogs. METHODS: We documented the timing of completed eruption of permanent dentition into occlusion (dental maturity) and the timing of growth plate closure at the proximal humerus (skeletal maturity) in ontogenetic series of wolves and 15 domestic dog breeds. Data for 137 domestic dog and 64 wolf individuals were collected based on radiographs and examination of macerated bones. RESULTS: Our analyses show that domestic dogs exhibit a similar sequence of dental and skeletal maturity as the ancestral wolf. Although the absolute change of the age at attainment of sexual maturity is great in domestic dogs as compared to the wolf, the sequence of dental, skeletal, and sexual maturity is not altered as extensively, contradicting one previous hypothesis. Moreover, our data suggest that the chondrodystrophic dachshund attains skeletal maturity earlier than the non-chondrodystrophic breeds examined here. CONCLUSIONS: Domestic dogs are more wolf-like in terms of the sequence of dental, skeletal, and sexual maturation than previously hypothesised. This implies that the domestication process and/or breed formation did not have a major impact on this sequence, although the absolute values of life history variables do have a greater range of variation than in the wild wolf.