Visual fields in hornbills: precision-grasping and sunshades

Retinal visual fields were determined in Southern Ground Hornbills Bucorvus leadbeateri and Southern Yellow-billed Hornbills Tockus leucomelas (Coraciiformes, Bucerotidae) using an ophthalmoscopic reflex technique. In both species the binocular field is relatively long and narrow with a maximum width of 30° occurring 40° above the bill. The bill tip projects into the lower half of the binocular field. This frontal visual field topography exhibits a number of key features that are also found in other terrestrial birds. This supports the hypothesis that avian visual fields are of three principal types that are correlated with the degree to which vision is employed when taking food items, rather than with phylogeny. However, unlike other species studied to date, in both hornbill species the bill intrudes into the binocular field. This intrusion of the bill restricts the width of the binocular field but allows the birds to view their own bill tips. It is suggested that this is associated with the precision-grasping feeding technique of hornbills. This involves forceps-like grasping and manipulation of items in the tips of the large decurved bill. The two hornbill species differ in the extent of the blind area perpendicularly above the head. Interspecific comparison shows that eye size and the width of the blind area above the head are significantly correlated. The limit of the upper visual field in hornbills is viewed through the long lash-like feathers of the upper lids and these appear to be used as a sunshade mechanism. In Ground Hornbills eye movements are non-conjugate and have sufficient amplitude (30–40°) to abolish the frontal binocular field and to produce markedly asymmetric visual field configurations.     

The subtlety of simple eyes: the tuning of visual fields to perceptual challenges in birds

Birds show interspecific variation both in the size of the fields of individual eyes and in the ways that these fields are brought together to produce the total visual field. Variation is found in the dimensions of all main parameters: binocular region, cyclopean field and blind areas. There is a phylogenetic signal with respect to maximum width of the binocular field in that passerine species have significantly broader field widths than non-passerines; broadest fields are found among crows (Corvidae). Among non-passerines, visual fields show considerable variation within families and even within some genera. It is argued that (i) the main drivers of differences in visual fields are associated with perceptual challenges that arise through different modes of foraging, and (ii) the primary function of binocularity in birds lies in the control of bill position rather than in the control of locomotion. The informational function of binocular vision does not lie in binocularity per se (two eyes receiving slightly different information simultaneously about the same objects from which higher-order depth information is extracted), but in the contralateral projection of the visual field of each eye. Contralateral projection ensures that each eye receives information from a symmetrically expanding optic flow-field from which direction of travel and time to contact targets can be extracted, particularly with respect to the control of bill position.     

Anatomical specializations for nocturnality in a critically endangered parrot, the Kakapo (Strigops habroptilus)

The shift from a diurnal to nocturnal lifestyle in vertebrates is generally associated with either enhanced visual sensitivity or a decreased reliance on vision. Within birds, most studies have focused on differences in the visual system across all birds with respect to nocturnality-diurnality. The critically endangered Kakapo (Strigops habroptilus), a parrot endemic to New Zealand, is an example of a species that has evolved a nocturnal lifestyle in an otherwise diurnal lineage, but nothing is known about its' visual system. Here, we provide a detailed morphological analysis of the orbits, brain, eye, and retina of the Kakapo and comparisons with other birds. Morphometric analyses revealed that the Kakapo's orbits are significantly more convergent than other parrots, suggesting an increased binocular overlap in the visual field. The Kakapo exhibits an eye shape that is consistent with other nocturnal birds, including owls and nightjars, but is also within the range of the diurnal parrots. With respect to the brain, the Kakapo has a significantly smaller optic nerve and tectofugal visual pathway. Specifically, the optic tectum, nucleus rotundus and entopallium were significantly reduced in relative size compared to other parrots. There was no apparent reduction to the thalamofugal visual pathway. Finally, the retinal morphology of the Kakapo is similar to that of both diurnal and nocturnal birds, suggesting a retina that is specialised for a crepuscular niche. Overall, this suggests that the Kakapo has enhanced light sensitivity, poor visual acuity and a larger binocular field than other parrots. We conclude that the Kakapo possesses a visual system unlike that of either strictly nocturnal or diurnal birds and therefore does not adhere to the traditional view of the evolution of nocturnality in birds.     

Binocular visual processing in the owl's telencephalon.

Single neurons recorded from the owl's visual Wulst are surprisingly similar to those found in mammalian striate cortex. The receptive fields of Wulst neurons are elaborated, in an apparently hierarchical fashion, from those of their monocular, concentrically organized inputs to produce binocular interneurons with increasingly sophisticated requirements for stimulus orientation, movement and binocular disparity. Output neurons located in the superficial laminae of the Wulst are the most sophisticated of all, with absolute requirements for a combination of stimuli, which include binocular presentation at a particular horizontal binocular disparity, and with no response unless all of the stimulus conditions are satisfied simultaneously. Such neurons have the properties required for 'global stereopsis', including a receptive field size many times larger than their optimal stimulus, which is more closely matched to the receptive fields of the simpler, disparity-selective interneurons. These marked similarities in functional organization between the avian and mammalian systems exist in spite of a number of structural differences which reflect their separate evoluntionary origins. Discussion therefore includes the possibility that there may exist for nervous systems only a very small number of possible solutions, perhaps a unique one, to the problem of stereopsis.     

The visual fields of Common Guillemots Uria aalge and Atlantic Puffins Fratercula arctica: foraging,vigilance and collision vulnerability

Significant differences in avian visual fields are found between closely related species that differ in their foraging technique. We report marked differences in the visual fields of two auk species. In air, Common Guillemots Uria aalge have relatively narrow binocular fields typical of those found in non‐passerine predatory birds. Atlantic Puffins Fratercula arctica have much broader binocular fields similar to those that have hitherto been recorded in passerines and in a penguin. In water, visual fields narrow considerably and binocularity in the direction of the bill is probably abolished in both auk species. Although perceptual challenges associated with foraging are similar in both species during the breeding season, when they are piscivorous, Puffins (but not Guillemots) face more exacting perceptual challenges when foraging at other times, when they take a high proportion of small invertebrate prey. Capturing this prey probably requires more accurate, visually guided bill placement and we argue that this is met by the Puffin's broader binocular field, which is retained upon immersion; its upward orientation may enable prey to be seen in silhouette. These visual field configurations have potentially important consequences that render these birds vulnerable to collision with human artefacts underwater, but not in air. They also have consequences for vigilance behaviour.     

Eye structure and foraging in King Penguins Aptenodytes patagonicus

Anterior eye structure and retinal visual fields were determined in King Penguins Aptenodytes patagonicus using keratometry and an ophthalmoscopic reflex technique. The cornea is relatively flat (radius 32.9 mm) and hence of low refractive power (10.2 dioptres in air) and this may be correlated with the amphibious nature of penguin vision. The large size of the eye and of the fully dilated pupil may be correlated with activity at low light levels. In air, the binocular field is long (vertical extent 180̀) and narrow (maximum width 29̀), with the bill placed approximately centrally—a topography found in a range of bird species which employ visual guidance of bill position when foraging. Upon immersion in water, the optical power of the cornea is abolished, with the effect that the monocular fields decrease and binocularity is lost. King Penguins have a pupil type which has not hitherto been recorded in birds. In daylight it contracts to a square-shaped pinhole but dilates to a large circular aperture in darkness. This change alters retinal illumination by 300-fold (2.5 log₁₀ units). When diving, this permits the retina to be pre-adapted to the low ambient light levels that the birds encounter upon reaching mesopelagic depths. These penguins also forage at depths where ambient light levels, even during the day, can fall below the equivalent of terrestrial starlight. Under these conditions, the birds must rely upon the detection of light from the photophores of their prey. In this they are aided by their absolutely large pupil size and broad cyclopean visual field.     

The visual fields of two ground‐foraging birds,House Finches and House Sparrows,allow for simultaneous foraging and anti‐predator vigilance

In birds, differences in the extent and position of the binocular visual field reflect adaptations to varying foraging strategies, and the extent of the lateral portion of the field may reflect anti‐predator strategies. The goal of this study was to describe and compare the visual fields of two ground‐foraging passerines, House Finch Carpodacus mexicanus and House Sparrow Passer domesticus. We found that both species have a binocular field type that is associated with the accurate control of bill position when pecking. Both species have eye movements of relatively large amplitude, which can produce substantial variations in the configuration of the binocular fields. We propose that in these ground foragers, their relatively wide binocular fields could function to increase foraging efficiency by locating multiple rather than single food items prior to pecking events. The lateral fields of both species are wide enough to facilitate the detection of predators or conspecifics while head‐down foraging. This suggests that foraging and scanning are not mutually exclusive activities in these species, as previously assumed. Furthermore, we found some slight, but significant, differences between species: House Sparrow binocular fields are both wider and vertically taller, and the blind area is wider than in House Finches. These differences may be related to variations in the degree of eye movements and position of the orbits in the skull.     

Hawk Eyes I: Diurnal Raptors Differ in Visual Fields and Degree of Eye Movement

Background

Different strategies to search and detect prey may place specific demands on sensory modalities. We studied visual field configuration, degree of eye movement, and orbit orientation in three diurnal raptors belonging to the Accipitridae and Falconidae families.

Methodology/Principal Findings

We used an ophthalmoscopic reflex technique and an integrated 3D digitizer system. We found inter-specific variation in visual field configuration and degree of eye movement, but not in orbit orientation. Red-tailed Hawks have relatively small binocular areas (∼33°) and wide blind areas (∼82°), but intermediate degree of eye movement (∼5°), which underscores the importance of lateral vision rather than binocular vision to scan for distant prey in open areas. Cooper''s Hawks'' have relatively wide binocular fields (∼36°), small blind areas (∼60°), and high degree of eye movement (∼8°), which may increase visual coverage and enhance prey detection in closed habitats. Additionally, we found that Cooper''s Hawks can visually inspect the items held in the tip of the bill, which may facilitate food handling. American Kestrels have intermediate-sized binocular and lateral areas that may be used in prey detection at different distances through stereopsis and motion parallax; whereas the low degree eye movement (∼1°) may help stabilize the image when hovering above prey before an attack.

Conclusions

We conclude that: (a) there are between-species differences in visual field configuration in these diurnal raptors; (b) these differences are consistent with prey searching strategies and degree of visual obstruction in the environment (e.g., open and closed habitats); (c) variations in the degree of eye movement between species appear associated with foraging strategies; and (d) the size of the binocular and blind areas in hawks can vary substantially due to eye movements. Inter-specific variation in visual fields and eye movements can influence behavioral strategies to visually search for and track prey while perching.     

The evolution of stereopsis and the Wulst in caprimulgiform birds: a comparative analysis

Owls possess stereopsis (i.e., the ability to perceive depth from retinal disparity cues), but its distribution amongst other birds has remained largely unexplored. Here, we present data on species variation in brain and telencephalon size and features of the Wulst, the neuroanatomical substrate that subserves stereopsis, in a putative sister-group to owls, the order Caprimulgiformes. The caprimulgiforms we examined included nightjars (Caprimulgidae), owlet-nightjars (Aegothelidae), potoos (Nyctibiidae), frogmouths (Podargidae) and the Oilbird (Steatornithidae). The owlet-nightjars and frogmouths shared almost identical relative brain, telencephalic and Wulst volumes as well as overall brain morphology and Wulst morphology with owls. Specifically, the owls, frogmouths and owlet-nightjars possess relatively large brains and telencephalic and Wulst volumes, had a characteristic brain shape and displayed prominent laminae in the Wulst. In contrast, potoos and nightjars both had relatively small brains and telencephala, and Wulst volumes that are typical for similarly sized birds from other orders. The Oilbird had a large brain, telencephalon and Wulst, although these measures were not quite as large as those of the owls. This gradation of owl-like versus nightjar-like brains within caprimulgiforms has significant implications for understanding the evolution of stereopsis and the Wulst both within the order and birds in general.     

Responses and properties of receptive fields of neurons in the visual projection zone of the pigeon hyperstriatum

Unit responses in the hyperstriatal region of the pigeon forebrain to the action of various visual stimuli were investigated. Particular attention was paid to the discovery of retinotopic projection in the Wulst region. It was shown that as the electrode was advanced in the caudal direction in the zone of visual projection of the hyperstriatum the receptive fields of the neurons recorded shifted in the opposite direction in the visual field. The receptive fields of neurons of the ventral and dorsal hyperstriatum lie higher in the visual field and are larger in diameter than those of neurons of the accessory hyperstriatum. Unit responses in the visual projection zone of the Wulst depend on the intensity of illumination, size, and speed and direction of movement of the test objects across the receptive field. The functional role of the retino-thalamo-telencephalic system in visual interpretation in birds is discussed and it is suggested that the Wulst region is comparable with the striatal and also with the frontal regions of the mammalian cortex.M. V. Lomonosov Moscow State University. Translated from Neirofiziologiya, Vol. 8, No. 3, pp. 230–236, May–June, 1976.     

Effects of activity pattern on eye size and orbital aperture size in primates

Among primates, nocturnal species exhibit relatively larger orbital apertures than diurnal species. Most researchers have considered this disparity in orbital aperture size to reflect differences in eye size, with nocturnal primates having relatively large eyes in order to maximize visual sensitivity. Presumed changes in eye size due to shifts in activity pattern are an integral part of theoretical explanations for many derived features of anthropoids, including highly convergent orbits and a postorbital septum. Here I show that despite clear differences in relative orbital aperture size, many diurnal and nocturnal primates do not differ in relative eye size. Among nocturnal primates, relative eye size is influenced by diet. Nocturnal visual predators (e.g., Tarsius, Loris, and Galago moholi) tend to have larger relative eye sizes than diurnal primates. By contrast, nocturnal frugivores (e.g., Perodicticus, Nycticebus, and Cheirogaleus) have relative eye sizes that are comparable to those of diurnal primates. Although some variation in orbital aperture size can be attributed to variation in eye size, both cornea size and orbit orientation also exert a strong influence on orbital aperture size. These findings argue for caution in the use of relative orbital aperture size as an indicator of activity pattern in fossil primates. These findings further suggest that existing scenarios for the evolution of unique orbital morphologies in anthropoids must be modified to reflect the importance of ecological variables other than activity pattern.     

Visual fields in the Black-crowned Night Heron Nycticorax nycticorax: nocturnality does not result in owl-like features

Compared with diurnally active species, the eyes of nocturnally active herons (Ardeidae) are relatively larger and more widely separated. These features are found in comparisons between the nocturnally foraging Black-crowned Night Heron Nycticorax nycticorax and the diurnally active Cattle Egret Bubulcus ibis. Casual viewing of the Black-crowned Night Heron gives the impression of a somewhat owl-like appearance, with an apparently wide degree of binocular overlap. Visual fields and eye movements were determined in two alert, restrained Black-crowned Night Herons with the use of an ophthalmoscopic technique. A wide degree of binocular overlap was not confirmed, and the Black-crowned Night Heron's visual field closely resembles those of diurnally foraging herons (Western Reef Heron Egretta gularis schistacea , Squacco Heron Ardeola ralloides , Cattle Egret). The Black-crowned Night Heron's binocular field is vertically long and narrow (maximum width 22̀), with the bill placed approximately at the centre. Monocular and cyclopean field widths in the horizontal plane equal 171̀ and 320̀, respectively. Retinal binocular overlap can be abolished by eye movements. There is a blind sector (10–13̀ wide) at the margin of each eye's optical field, and this results in the functional retinal binocular field being much narrower than the optical binocular fields. It is these blind sectors which give rise to the appearance of a much wider binocular field. The visual field characteristics of this heron species may be best understood in relation to the foraging technique of capturing agile, evasive prey directly in the bill. The comparatively large size of the Black-crowned Night Herons' eyes may be associated with activity over a wide range of natural light levels but does not give rise to binocular fields larger than those of diurnal heron species.     

White‐headed Vulture Trigonoceps occipitalis shows visual field characteristics of hunting raptors

The visual fields of the Aegypiinae vultures have been shown to be adapted primarily to meet two key perceptual challenges of their obligate carrion‐feeding behaviour: scanning the ground and preventing the sun's image falling upon the retina. However, field observations have shown that foraging White‐headed Vultures Trigonoceps occipitalis are not exclusively carrion‐feeders; they are also facultative predators of live prey. Such feeding is likely to present perceptual challenges that are additional to those posed by carrion‐feeding. Binocularity is the key component of all visual fields and in birds it is thought to function primarily in the accurate placement and time of contact of the talons and bill, especially in the location and seizure of food items. We determined visual fields in White‐headed Vultures and compared them with those of two species of carrion‐eating Gyps vultures. The visual field of White‐headed Vultures has more similarities with those of predatory raptors (e.g. accipitrid hawks) than with the taxonomically more closely related Gyps vultures. Maximum binocular field width in White‐headed Vultures (30°) is significantly wider than that in Gyps vultures (20°). The broader binocular fields in White‐headed Vultures probably facilitate accurate placement and timing of the talons when capturing evasive live prey.     

The eye of the humboldt penguin, Spheniscus humboldti: visual fields and schematic optics

Construction of a schematic eye indicates that the eye of Spheniscus humboldti is aquatic in design. The lens has a power of 100 dioptres (D) while (in air) the cornea has a power of 29 D. In air, the eye is myopic (approximately 28 D) but in water it is emmetropic. Minimum pupil size would seem insufficient to allow the pupil to function as a stenopaic aperture and increase depth of focus sufficiently to overcome the eye's aerial myopia. Entry into water reduces maximum image brightness by approximately three times. In air, the maximum width of the retinal binocular field is 45 degrees and this occurs approximately 10 degrees above the line of the bill. The bill intrudes into the retinal field and binocular field width in the plane containing the bill and the optic axes is 28 degrees. The vertical extent of the binocular field is 125 degrees. In the plane containing the optic axes the cyclopean field equals 282 degrees and the optic axes diverge by 116 degrees. In this plane the mean uniocular field is 155 degrees with the temporal hemifield approximately 11 degrees larger than the nasal hemifield. Entry into water reduces the widths of the visual fields such that maximum binocular field width is only 17 degrees and the vertical extent is reduced to about 80 degrees. Binocular vision is lost in the plane of the bill, and the uniocular retinal field is reduced by 32 degrees and the cyclopean field by 36 degrees.     

Visual fields in woodcocks Scolopax rusticola (Scolopacidae; Charadriiformes)

Woodcocks, Scolopax rusticola, are long-billed terrestrial wading birds (Scolopacidae; Charadriiformes) which forage primarily by probing in soft substrates for invertebrates. Visual field topography in restrained alert birds was investigated using an ophthalmoscopic reflex technique.

1. Eye movements of significant amplitude are absent.
2. The retinal binocular field is long and narrow. It extends through 190° in the median sagittal plane. When the head adopts a normal posture (bill at an angle of 40° below the horizontal) the binocular field stretches from 25° above the bill to 5° above the horizontal behind the head. Thus, woodcocks have comprehensive visual coverage of the hemisphere above them but the bill falls outside the visual field. Maximum binocular field width equals 12° and occurs perpendicular to the line of the bill. To the rear of the head binocular field width is less than 5° except in an area 40° above the horizontal where it increases to 7°.
3. Monocular retinal fields in the horizontal plane are 182° wide. There is no blind sector at the margin of the optical fields.
4. The general structure of woodcock skulls facilitates panoramic vision in a horizontal plane.
5. Interspecific comparisons are consistent with the hypothesis that visual field topography among birds is closely associated with the role of vision in foraging. Comprehensive visual coverage of the celestial hemisphere probably occurs only in species, such as woodcocks, which rely primarily upon senses other than vision to guide foraging.

     

Vision and the foraging technique of Great Cormorants Phalacrocorax carbo: pursuit or close‐quarter foraging?

Predatory diving birds, such as cormorants (Phalacrocoracidae), have been generally regarded as visually guided pursuit foragers. However, due to their poor visual resolution underwater, it has recently been hypothesized that Great Cormorants do not in fact employ a pursuit-dive foraging technique. They appear capable of detecting typical prey only at short distances, and primarily use a foraging technique in which prey may be detected only at close quarters or flushed from a substratum or hiding place. In birds, visual field parameters, such as the position and extent of the region of binocular vision, and how these are altered by eye movements, appear to be determined primarily by feeding ecology. Therefore, to understand further the feeding technique of Great Cormorants we have determined retinal visual fields and eye movement amplitudes using an ophthalmoscopic reflex technique. We show that visual fields and eye movements in cormorants exhibit close similarity with those of other birds, such as herons (Ardeidae) and hornbills (Bucerotidae), which forage terrestrially typically using a close-quarter prey detection or flushing technique and/or which need to examine items held in the bill before ingestion. We argue that this visual field topography and associated eye movements is a general characteristic of birds whose foraging requires the detection of nearby mobile prey items from within a wide arc around the head, accurate capture of that prey using the bill, and visual examination of the caught prey held in the bill. This supports the idea that cormorants, although visually guided predators, are not primarily pursuit predators, and that their visual fields exhibit convergence towards a set of characteristics that meet the perceptual challenges of close-quarter prey detection or flush foraging in both aquatic and terrestrial environments.     

Eye shape and activity pattern in birds

Many aspects of an animal's ecology are associated with activity pattern, the time of day when that animal is awake and active. There are two major activity patterns: diurnal , active during the day in a light-rich, or photopic, environment, and nocturnal , active after sunset in a light-limited, or scotopic, environment. Birds are also cathemeral , or equally likely to be awake at any time of day, or crepuscular , awake and active at dawn and dusk. Each of these activity patterns is associated with different levels of ambient light. This study examines how the morphology (size and shape) of the eye varies according to these different light environments for birds in a phylogenetic context. Activity pattern has a significant influence on eye shape and size in birds. Birds that are adapted for scotopic vision have eye shapes that are optimized for visual sensitivity, with larger corneal diameters relative to axial lengths. Birds that are adapted for photopic vision have eye shapes that are optimized for visual acuity, with larger axial lengths relative to corneal diameters. Birds adapted for scotopic vision also exhibit absolutely larger corneal diameters and axial lengths than do photopic birds. The results indicate that the light level under which the bird functions has a more significant influence on eye shape than phylogeny.     

Spontaneous retinal activity mediates development of ocular dominance columns and binocular receptive fields in v1

The mechanisms that give rise to ocular dominance columns (ODCs) during development are controversial. Early experiments indicated a key role for retinal activity in ODC formation. However, later studies showed that in those early experiments, the retinal activity perturbation was initiated after ODCs had already formed. Moreover, recent studies concluded that early eye removals do not impact ODC segregation. Here we blocked spontaneous retinal activity during the very early stages of ODC development. This permanently disrupted the anatomical organization of ODCs and led to a dramatic increase in receptive field size for binocular cells in primary visual cortex. Our data suggest that early spontaneous retinal activity conveys crucial information about whether thalamocortical axons represent one or the other eye and that this activity mediates binocular competition important for shaping receptive fields in primary visual cortex.     

Shifted magnetic fields lead to deflected and axial orientation of migrating robins,Erithacus rubecula,at sunset

The migratory orientation of the robin was tested in shifted magnetic fields during the twilight period after sunset, under clear skies and under simulated total overcast. The horizontal direction of the geomagnetic field was shifted 90° to the right or left in relation to the local magnetic field, without changing either the intensity of the field or its angle of inclination. Experiments were conducted during both spring and autumn, with robins captured as passage migrants at the Falsterbo and Ottenby bird observatories in southern Sweden as test subjects. Generally, the orientation of robins was affected by magnetic shifts compared to controls tested in the natural geomagnetic field. Autumn birds from the two capture sites differed in their responses, probably because of different migratory dispositions and body conditions. The robins most often changed their orientation to maintain their typical axis of migration relative to the shifted magnetic fields. However, preferred directions in relation to the shifted magnetic fields were frequently reverse from normal, or axial rather than unimodal. These results disagree with suggested mechanisms for orientation by visual sunset cues and with the proposed basis of magnetic orientation. They do, however, demonstrate that the geomagnetic field is involved in the sunset orientation of robins, probably in combination with additional visual or non-visual cues that contribute to establish magnetic polarity.     

Function of the mammalian postorbital bar

Complete postorbital bars, bony arches that encompass the lateral aspect of the eye and form part of a circular orbit, have evolved homoplastically multiple times during mammalian evolution. Numerous functional hypotheses have been advanced for postorbital bars, the most promising being that postorbital bars function to stiffen the lateral orbit in taxa that have significant angular deviation between the temporal fossa and the bony orbit. Without a stiff lateral orbit the anterior temporalis muscle and fascia potentially would pull on the postorbital ligament, deform the orbit, and cause disruption of oculomotor precision. Morphometric data were collected on 1,329 specimens of 324 taxa from 16 orders of extant eutherian and metatherian mammals in order to test whether the orientation of the orbit relative to the temporal fossa is correlated with the replacement of the postorbital ligament with bone. The allometric and ecological influences on orbit orientation across mammals are also explored. The morphometric results corroborate the hypothesis: Shifts in orbit orientation relative to the temporal fossa are correlated with the size of the postorbital processes, which replace the ligament. The allometric and ecological factors that influence orbit orientation vary across taxa. Postorbital bars stiffen the lateral orbital wall. Muscle pulleys, ligaments, and other connective tissue attach to the lateral orbital wall, including the postorbital bar. Without a stiff lateral orbit, deformation due to temporalis contraction would displace soft tissues contributing to normal oculomotor function.