The evolutionary origin of avian facial bristles and the likely role of rictal bristles in feeding ecology.

Facial bristles are one of the least described feather types and have not yet been systematically studied across phylogenetically diverse avian species. Consequently, little is known about their form, function and evolutionary history. Here we address this knowledge gap by characterising the evolution of facial bristles for the first time. We especially focus on rictal bristle presence and their associations with foraging behaviour, diet and habitat preferences in 1022 avian species, representing 91 families in 29 orders. Results reveal that upper rictal, lower rictal and interramal bristles were likely to be present in the most recent common ancestor of this avian phylogeny, whereas narial bristles were likely to be absent. Rictal bristle presence, length and shape varied both within and between avian orders, families and genera. Rictal bristles were gained or lost multiple times throughout evolution, which suggest that the different morphologies observed within species might not be homologous. Phylogenetic relatedness is also not likely to be the only driver of rictal bristle presence and morphology. Rictal bristle presence and length were associated with species-specific ecological traits, especially nocturnality. Our findings suggest that species foraging in low-light conditions are likely to have longer rictal bristles, and that rictal bristles are likely to have evolved in early birds.

Anatomy of bristles on the nares and rictus of western barn owls (Tyto alba).

Many nocturnal avian species, such as Strigiformes, Caprimulgiformes and Apterygiformes, have sensitive vibrotactile bristles on their upper bill, especially on their rictus. The anatomy of these bristles can vary, especially in terms of sensitivity (Herbst corpuscle number), bristle length and bristle number. This variation is thought to be associated with foraging - such that diurnal, open foragers have smaller and less-sensitive bristles. Here, we describe bristle morphology and follicle anatomy in the western barn owl (Tyto alba) for the first time, using both live and roadkill wild owls. We show that T. alba have both narial and rictal bristles that are likely to be vibrotactile, since they have Herbst corpuscles around their follicles. We observed more numerous (~8) and longer bristles (~16 mm) on the nares of T. alba, than on the rictal region (~4 and ~13 mm respectively). However, the narial bristle follicles contained fewer Herbst corpuscles in their surroundings (~5) than the rictal bristles (~7); indicating that bristle length is not indicative of sensitivity. As well as bristle length and number varying between different facial regions, they also varied between individuals, although the cause of this variation remains unclear. Despite this variation, the gross anatomy of facial bristle follicles appears to be conserved between nocturnal Strigiformes, Caprimulgiformes and Apterygiformes. Understanding more about how T. alba use their bristles would, therefore, give us greater insights into the function of avian bristles in general.

Anatomy of avian rictal bristles in Caprimulgiformes reveals reduced tactile function in open-habitat, partially diurnal foraging species.

Avian rictal bristles are present in many species of birds, especially in nocturnal species. Rictal bristles occur along the upper beak and are morphologically similar to mammalian whiskers. Mammalian whiskers are important tactile sensors, guiding locomotion, foraging and social interactions, and have a well-characterised anatomy. However, it is not yet known whether avian rictal bristles have a sensory function, and their morphology, anatomy and function have also not been described in many species. Our study compares bristle morphology, follicle anatomy and their association with foraging traits, across 12 Caprimulgiform species. Rictal bristle morphology and follicle anatomy were diverse across the 12 species. Nine of the 12 species had mechanoreceptors around their bristle follicles; however, there was large variation in their musculature, mechanoreceptor numbers and bristle morphology. Overall, species with short, thin, branching bristles that lacked mechanoreceptors tended to forage pre-dusk in open habitats, whereas species with mechanoreceptors around their bristle follicle tended to forage at night and in more closed habitats. We suggest that rictal bristles are likely to be tactile in many species and may aid in navigation, foraging and collision avoidance; however, identifying rictal bristle function is challenging and demands further investigation in many species.

Paw Morphology in the Domestic Guinea Pig (Cavia porcellus) and Brown Rat (Rattus norvegicus).

Mammals have adapted to different habitats, food types, and modes of locomotion, which are reflected in a diverse range of paw morphologies. While the behavior of rats and guinea pigs is well defined, especially in terms of their locomotor and foraging behaviors, the anatomy of their foot pads has not yet been explored and compared. This study investigated adaptations in paw morphology in the domestic guinea pig (Cavia porcellus) and the brown rat (Rattus norvegicus). We predicted that rat paws would display adaptations associated with paw dexterity for handling prey items and climbing, whereas guinea pig paws would support mechanical pressure absorption for a herbivorous, sedentary, and terrestrial lifestyle. Using histology techniques and scanning electron microscope, we show that rat paws have many small, deformable pads that are relatively spaced out to enable movement. The pads are clustered toward the anterior of the foot, which coincides with where the most force occurs during locomotion, as rats walk on their toes and toward the front of their paw. Guinea pigs had fewer and larger pads and the posterior pad of the forepaw was textured and contained cartilage, which may act to reduce friction and compression during standing and locomotion. We suggest that differences in paw morphology in rat and guinea pig are associated with loading during locomotion as well as paw mobility. Examining paw morphology and movement abilities in more species will give further insights in to the evolution of locomotor adaptations and paw dexterity in rodents. Anat Rec, 302:2300-2310, 2019. © 2019 American Association for Anatomy.

Mystacial Whisker Layout and Musculature in the Guinea Pig (Cavia porcellus): A Social, Diurnal Mammal.

All mammals (apart from apes and humans) have whiskers that make use of a similar muscle arrangement. Whisker specialists, such as rats and mice, tend to be nocturnal and arboreal, relying on their whisker sense of touch to guide exploration around tree canopies at night. As such, nocturnal arboreal rodents have many whiskers that are organised into a grid-like pattern, and moved using a complex array of muscles. Indeed, most arboreal, nocturnal mammals tend to have specialised whiskers that are longer and arranged in a dense, regular grid, compared with terrestrial, diurnal mammals. The guinea pig diverged early from murid rodents (around 75 million years ago), and are ground-dwelling, diurnal animals. It would be predicted that, as a terrestrial mammal, they may have less whiskers and a reduced muscle architecture compared to arboreal, nocturnal rodents. We examined the mystacial whisker layout, musculature and movement capacity of Guinea pig (Cavia porcellus) whiskers and found that they did indeed have a disorganized whisker layout, with a fortification around the eye area. In addition, there was a reduction in musculature, especially in the intrinsic muscles. Despite guinea pigs not cyclically moving their whiskers, the mystacial musculature was still very similar to that of murid rodents. We suggest that the conserved presence of whisker layout and musculature, even in visual mammals such as primates and guinea pigs, may indicate that whiskers still play an important role in these animals, including protecting the eyes and being involved in tactile social behaviors. Anat Rec, 300:527-536, 2017. © 2016 Wiley Periodicals, Inc.