The multifactorial nature of beak and skull shape evolution in parrots and cockatoos (Psittaciformes).

BACKGROUND: The Psittaciformes (parrots and cockatoos) are characterised by their large beaks, and are renowned for their ability to produce high bite forces. These birds also possess a suite of modifications to their cranial architecture interpreted to be adaptations for feeding on mechanically resistant foods, yet the relationship between cranial morphology and diet has never been explicitly tested. Here, we provide a three-dimensional geometric morphometric analysis of the developmental and biomechanical factors that may be influencing the evolution of psittaciformes' distinctive cranial morphologies. RESULTS: Contrary to our own predictions, we find that dietary preferences for more- or less- mechanically resistant foods have very little influence on beak and skull shape, and that diet predicts only 2.4% of the shape variation in psittaciform beaks and skulls. Conversely, evolutionary allometry and integration together predict almost half the observed shape variation, with phylogeny remaining an important factor in shape identity throughout our analyses, particularly in separating cockatoos (Cacatuoidea) from the true parrots (Psittacoidea). CONCLUSIONS: Our results are similar to recent findings about the evolutionary trajectories of skull and beak shape in other avian families. We therefore propose that allometry and integration are important factors causing canalization of the avian head, and while diet clearly has an influence on beak shape between families, this may not be as important at driving evolvability within families as is commonly assumed.

New Insight into Parrots' Mitogenomes Indicates That Their Ancestor Contained a Duplicated Region.

Mitochondrial genomes of vertebrates are generally thought to evolve under strong selection for size reduction and gene order conservation. Therefore, a growing number of mitogenomes with duplicated regions changes our view on the genome evolution. Among Aves, order Psittaciformes (parrots) is especially noteworthy because of its large morphological, ecological, and taxonomical diversity, which offers an opportunity to study genome evolution in various aspects. Former analyses showed that tandem duplications comprising the control region with adjacent genes are restricted to several lineages in which the duplication occurred independently. However, using an appropriate polymerase chain reaction strategy, we demonstrate that early diverged parrot groups contain mitogenomes with the duplicated region. These findings together with mapping duplication data from other mitogenomes onto parrot phylogeny indicate that the duplication was an ancestral state for Psittaciformes. The state was inherited by main parrot groups and was lost several times in some lineages. The duplicated regions were subjected to concerted evolution with a frequency higher than the rate of speciation. The duplicated control regions may provide a selective advantage due to a more efficient initiation of replication or transcription and a larger number of replicating genomes per organelle, which may lead to a more effective energy production by mitochondria. The mitogenomic duplications were associated with phenotypic features and parrots with the duplicated region can live longer, show larger body mass as well as predispositions to a more active flight. The results have wider implications on the presence of duplications and their evolution in mitogenomes of other avian groups.

Comparison of Dorsoventral Erect and Ventrodorsal Supine Radiographic Views for the Evaluation of Intracoelomic Organs in Clinically Normal African Grey Parrots (Psittacus erithacus).

Standard positioning for radiographic evaluation may require sedation and can be deleterious in critically ill birds. A prospective crossover study was performed in 15 clinically normal African grey parrots (Psittacus erithacus) to describe an alternative, unrestrained radiographic view. Whole-body radiographs were obtained in unrestrained dorsoventral erect (DVE) views and in anesthetized ventrodorsal supine (VDS) views. Visualization of various anatomic items in each view was scored by 3 observers. The surface area of the air sacs and the width of the heart, liver, thorax, and cardiohepatic waist were measured in DVE and VDS views. Measurements were obtained by 3 observers, and 1 observer repeated the measurements twice. Intraobserver and interobserver agreement were assessed. Major rotation of the coelom and superimposition of the limbs over the coelom were, respectively, observed in 4 of 15 (27%) and 15 of 15 (100%) of the DVE views and not observed in VDS views. The evaluation of the respiratory tract structures was considered limited in DVE views compared with VDS views, and the surface areas of the air sacs were significantly smaller. The proventriculus and ventriculus were more visible in the DVE view. The visualization of the heart was not significantly different between the 2 views. The absolute measures of heart, liver, and cardiohepatic waist width were significantly larger in the DVE view compared with the VDS view. Moderate intraobserver and interobserver agreement was observed in the evaluation of the 2 views. In conclusion, the DVE view could be adequate to assess the heart and the upper digestive tract. This positioning is likely to provide clinically relevant information for cases in which general anesthesia or dorsal recumbency is contraindicated.

Anatomy of the forelimb musculature and ligaments of Psittacus erithacus (Aves: Psittaciformes).

Parrots (order Psittaciformes) are a rather homogeneous group of birds that can be easily distinguished by the notably modified morphology of the skull and hindlimb. Detailed description of the forelimb morphology in these birds has never been provided, though parrots are often used as model objects in flight studies. Parrots are also considered the closest living relatives of the perching birds (Passeriformes), and thus knowledge of the wing morphology in Psittaciformes is important for understanding the evolution of the locomotor apparatus on the way to the most speciose group of birds. Here we provide a comprehensive illustrated description of the wing morphology (musculature and ligaments) of the African grey parrot (Psittacus erithacus) and compare it with several closely related taxa of the high clade Eufalconimorphae and more distantly related outgroups (based on personal dissections and literature data). We note a general similarity of the wing musculature between P. erithacus and Falconidae. A number of features common with the outgroup Columbidae supports a generally plesiomorphic structure of the forelimb in parrots as compared with the Passeriformes. Nevertheless, the wing of the Psittaciformes displays a series of structural (likely autapomorphic) modifications, which can be explained in terms of adaptations for flight with vertical body. An analysis of the anatomical data for parrots (ratio of wing elevators and highly unusual development of the M. supracoracoideus), which is based on the current experiment-based knowledge of the flapping flight in birds, allows us to hypothesize that parrots are able to produce useful aerodynamic force during the upstroke, which is also known for pigeons and hummingbirds. This supposed ability of vertical flight and the zygodactyl foot together link the origin of parrots with the dense (likely tropical) forests.

Comparative brain morphology of Neotropical parrots (Aves, Psittaciformes) inferred from virtual 3D endocasts.

Psittaciformes are a very diverse group of non-passerine birds, with advanced cognitive abilities and highly developed locomotor and feeding behaviours. Using computed tomography and three-dimensional (3D) visualization software, the endocasts of 14 extant Neotropical parrots were reconstructed, with the aim of analysing, comparing and exploring the morphology of the brain within the clade. A 3D geomorphometric analysis was performed, and the encephalization quotient (EQ) was calculated. Brain morphology character states were traced onto a Psittaciformes tree in order to facilitate interpretation of morphological traits in a phylogenetic context. Our results indicate that: (i) there are two conspicuously distinct brain morphologies, one considered walnut type (quadrangular and wider than long) and the other rounded (narrower and rostrally tapered); (ii) Psittaciformes possess a noticeable notch between hemisphaeria that divides the bulbus olfactorius; (iii) the plesiomorphic and most frequently observed characteristics of Neotropical parrots are a rostrally tapered telencephalon in dorsal view, distinctly enlarged dorsal expansion of the eminentia sagittalis and conspicuous fissura mediana; (iv) there is a positive correlation between body mass and brain volume; (v) psittacids are characterized by high EQ values that suggest high brain volumes in relation to their body masses; and (vi) the endocranial morphology of the Psittaciformes as a whole is distinctive relative to other birds. This new knowledge of brain morphology offers much potential for further insight in paleoneurological, phylogenetic and evolutionary studies.

The first fossil parrot (Aves, Psittaciformes) from Siberia and its implications for the historical biogeography of Psittaciformes.

Modern parrots (crown Psittaciformes) are a species-rich group of mostly tropical and subtropical birds with a very limited fossil record. A partial tarsometatarsus from the late Early Miocene of Siberia (Baikal Lake) is the first pre-Quaternary find of crown Psittaciformes in Asia (and Siberia in particular) and is also the northern-most find of this bird order worldwide. This find documents a broad geographical distribution of parrots during the warmest phase of the Miocene (the so-called 'Miocene Climatic Optimum'), which has implications for the historical biogeography of Psittaciformes. The presence of parrots on both sides of the Pacific Ocean at the end of the Early Miocene implies a (most probably eastwards) trans-Beringian dispersal which likely took place about 16-18 Ma. The broad Eurasian distribution of parrots in the past further supports a hypothesis that ancestors of modern genera Coracopsis and Agapornis could reach Africa from Eurasia.

Normal computed tomographic features and reference values for the coelomic cavity in pet parrots.

BACKGROUND: The increasing popularity gained by pet birds over recent decades has highlighted the role of avian medicine and surgery in the global veterinary scenario; such a need for speciality avian medical practice reflects the rising expectation for high-standard diagnostic imaging procedures. The aim of this study is to provide an atlas of matched anatomical cross-sections and contrast-enhanced CT images of the coelomic cavity in three highly diffused psittacine species. RESULTS: Contrast-enhanced computed tomographic studies of the coelomic cavity were performed in 5 blue-and-gold macaws, 4 African grey parrots and 6 monk parakeets by means of a 4-multidetector-row CT scanner. Both pre- and post-contrast scans were acquired. Anatomical reference cross-sections were obtained from 5 blue-and-gold macaw, 7 African grey parrot, and 9 monk parakeet cadavers. The specimens were stored in a -20 °C freezer until completely frozen and then sliced at 5-mm intervals by means of a band saw. All the slices were photographed on both sides. Individual anatomical structures were identified by means of the available literature. Pre- and post-contrast attenuation reference values for the main coelomic organs are reported in Hounsfield units (HU). CONCLUSIONS: The results provide an atlas of matched anatomical cross-sections and contrast-enhanced CT images of the coelomic cavity in three highly diffused psittacine species.

Amorphous diamond-structured photonic crystal in the feather barbs of the scarlet macaw.

Noniridescent coloration by the spongy keratin in parrot feather barbs has fascinated scientists. Nonetheless, its ultimate origin remains as yet unanswered, and a quantitative structural and optical description is still lacking. Here we report on structural and optical characterizations and numerical simulations of the blue feather barbs of the scarlet macaw. We found that the sponge in the feather barbs is an amorphous diamond-structured photonic crystal with only short-range order. It possesses an isotropic photonic pseudogap that is ultimately responsible for the brilliant noniridescent coloration. We further unravel an ingenious structural optimization for attaining maximum coloration apparently resulting from natural evolution. Upon increasing the material refractive index above the level provided by nature, there is an interesting transition from a photonic pseudogap to a complete bandgap.

Theoretical morphology and development of flight feather vane asymmetry with experimental tests in parrots.

Asymmetry in flight feather vane width is a major functional innovation associated with the evolution of flight in the ancestors of birds. However, the developmental and morphological basis of feather shape is not simple, and the developmental processes involved in vane width asymmetry are poorly understood. We present a theoretical model of feather morphology and development that describes the possible ways to modify feather development and produce vane asymmetry. Our model finds that the theoretical morphospace of feather shape is redundant, and that many different combinations of parameters could be responsible for vane asymmetry in a given feather. Next, we empirically measured morphological and developmental model parameters in asymmetric and symmetric feathers from two species of parrots to identify which combinations of parameters create vane asymmetry in real feathers. We found that both longer barbs, and larger barb angles in the relatively wider trailing vane drove asymmetry in tail feathers. Developmentally, longer barbs were the result of an offset of the radial position of the new barb locus, whereas larger barb angles were produced by differential expansion of barbs as the feather unfurls from the tubular feather germ. In contrast, the helical angle of barb ridge development did not contribute to vane asymmetry and could be indicative of a constraint. This research provides the first comprehensive description of both the morphological and developmental modifications responsible for vane asymmetry within real feathers, and identifies key steps that must have occurred during the evolution of vane asymmetry.

Retinal ganglion cell topography and spatial resolution of two parrot species: budgerigar (Melopsittacus undulatus) and Bourke's parrot (Neopsephotus bourkii).

Retinal ganglion cell (RGC) isodensity maps indicate important regions in an animal's visual field. These maps can also be combined with measures of focal length to estimate the theoretical visual acuity. Here we present the RGC isodensity maps and anatomical spatial resolving power in three budgerigars (Melopsittacus undulatus) and two Bourke's parrots (Neopsephotus bourkii). Because RGCs were stacked in several layers, we modified the Nissl staining procedure to assess the cell number in the whole-mounted and cross-sectioned tissue of the same retinal specimen. The retinal topography showed surprising variation; however, both parrot species had an area centralis without discernable fovea. Budgerigars also had a putative area nasalis never reported in birds before. The peak RGC density was 22,300-34,200 cells/mm(2) in budgerigars and 18,100-38,000 cells/mm(2) in Bourke's parrots. The maximum visual acuity based on RGCs and focal length was 6.9 cyc/deg in budgerigars and 9.2 cyc/deg in Bourke's parrots. These results are lower than earlier behavioural estimates. Our findings illustrate that retinal topography is not a very fixed trait and that theoretical visual acuity estimations based on RGC density can be lower than the behavioural performance of the bird.

Molecular and cellular changes associated with the evolution of novel jaw muscles in parrots.

Vertebrates have achieved great evolutionary success due in large part to the anatomical diversification of their jaw complex, which allows them to inhabit almost every ecological niche. While many studies have focused on mechanisms that pattern the jaw skeleton, much remains to be understood about the origins of novelty and diversity in the closely associated musculature. To address this issue, we focused on parrots, which have acquired two anatomically unique jaw muscles: the ethmomandibular and the pseudomasseter. In parrot embryos, we observe distinct and highly derived expression patterns for Scx, Bmp4, Tgfß2 and Six2 in neural crest-derived mesenchyme destined to form jaw muscle connective tissues. Furthermore, immunohistochemical analysis reveals that cell proliferation is more active in the cells within the jaw muscle than in surrounding connective tissue cells. This biased and differentially regulated mode of cell proliferation in cranial musculoskeletal tissues may allow these unusual jaw muscles to extend towards their new attachment sites. We conclude that the alteration of neural crest-derived connective tissue distribution during development may underlie the spatial changes in jaw musculoskeletal architecture found only in parrots. Thus, parrots provide valuable insights into molecular and cellular mechanisms that may generate evolutionary novelties with functionally adaptive significance.

Vision, touch and object manipulation in Senegal parrots Poicephalus senegalus.

Parrots are exceptional among birds for their high levels of exploratory behaviour and manipulatory abilities. It has been argued that foraging method is the prime determinant of a bird's visual field configuration. However, here we argue that the topography of visual fields in parrots is related to their playful dexterity, unique anatomy and particularly the tactile information that is gained through their bill tip organ during object manipulation. We measured the visual fields of Senegal parrots Poicephalus senegalus using the ophthalmoscopic reflex technique and also report some preliminary observations on the bill tip organ in this species. We found that the visual fields of Senegal parrots are unlike those described hitherto in any other bird species, with both a relatively broad frontal binocular field and a near comprehensive field of view around the head. The behavioural implications are discussed and we consider how extractive foraging and object exploration, mediated in part by tactile cues from the bill, has led to the absence of visual coverage of the region below the bill in favour of more comprehensive visual coverage above the head.

Colourful parrot feathers resist bacterial degradation.

The brilliant red, orange and yellow colours of parrot feathers are the product of psittacofulvins, which are synthetic pigments known only from parrots. Recent evidence suggests that some pigments in bird feathers function not just as colour generators, but also preserve plumage integrity by increasing the resistance of feather keratin to bacterial degradation. We exposed a variety of colourful parrot feathers to feather-degrading Bacillus licheniformis and found that feathers with red psittacofulvins degraded at about the same rate as those with melanin and more slowly than white feathers, which lack pigments. Blue feathers, in which colour is based on the microstructural arrangement of keratin, air and melanin granules, and green feathers, which combine structural blue with yellow psittacofulvins, degraded at a rate similar to that of red and black feathers. These differences in resistance to bacterial degradation of differently coloured feathers suggest that colour patterns within the Psittaciformes may have evolved to resist bacterial degradation, in addition to their role in communication and camouflage.

Scanning electron microscopic study of the tongue in the rainbow lorikeet (Trichoglossus haematodus).

The dorsal lingual surfaces of rainbow lorikeet (Trichoglossus haematodus) were examined by scanning electron microscopy. Macroscopically, the tongue of the rainbow lorikeet has a finger-like shape. Three parts are distinguished in the dorsal surface of the tongue: the apex, body, and root of the tongue. The apex of the tongue has numerous processes inclined toward medial side from lateral side. These processes are rod-like structure and smooth surfaces. Many grooves are observed in both lateral sides of the lingual body. A large opening of the lingual gland exists in central part of the lingual root and some large openings of the lingual glands exist in both lateral sides of the lingual root.

Radiographic measurement of internal organs in Spix's macaws (Cyanopsitta spixii).

Radiology is an important diagnostic instrument in avian medicine, but standard measurement ranges for the objective evaluation of radiographs of birds are rare. To establish radiographic reference ranges for the critically endangered Spix's macaw (Cyanopsitta spixii), we measured radiographic silhouettes of the heart, liver, kidneys, spleen, proventriculus, and keel of the sternum on 29 radiographs taken under standardized conditions in adult and juvenile, clinically healthy birds. Ratios were determined for the proventricular diameter-to-keel height, the width of the heart to the width of the thorax, and for the "hourglass shape" (ratio of the width of the heart to the width of the liver). No significant differences were found between the sexes among the adult birds. Compared with adult birds, juvenile females had a significantly larger heart width (19.8 +/- 1.4 mm versus 21.2 +/- 0.7 mm), ratio of the heart width to the thorax width (0.86 +/- 0.08 versus 0.94 +/- 0.09), and horizontal width of the spleen (7.7 +/- 0.6 mm versus 8.5 +/- 0.4 mm). Results of radiographic measurements in the Spix's macaws were comparable to those published from other psittacine species. These reference ranges will facilitate a more objective radiographic evaluation of captive Spix's macaws.

Study of laser actions by bird's feathers with photonic crystals.

Random lasers had been made by some biomaterials as light scattering materials, but natural photonic crystals have been rarely reported as scattering materials. Here we demonstrate the ability of natural photonic crystals to drive laser actions by sandwiched the feathers of the Turquoise-Fronted Amazon parrot and dye between two plastic films. Parrot feathers comprise abundant photonic crystals, and different color feathers compose of different ratios of the photonic crystal, which directly affect the feather reflectance. In this study, the multi-reflection scattering that occurred at the interface between the photonic crystal and gain media efficiently reduce the threshold; therefore, the more photonic crystal constitutes in the feathers; the lower threshold can be obtained. The random lasers can be easily made by the integration of bird feather photonic crystals and dye with a simple and sustainable manufacturing approach.

Respiratory Diseases of Parrots: Anatomy, Physiology, Diagnosis and Treatment.

This article is aimed to help the reader to understand better how to diagnose and treat different respiratory diseases in Psittaciformes (parrot-like birds). The article starts from a review of avian respiratory anatomy and physiology, and then moves forward into diagnostic techniques, most common diseases, split in species and anatomic location, and common treatment regimens.

Computed tomographic and radiographic morphometric study of cardiac and coelomic dimensions in captive blue-fronted Amazon parrots (Amazona aestiva, Linnaeus, 1758) with varying body condition scores.

This study aimed to assess radiographic and tomographic cardiac parameters, including width and length of the heart, and the ratio of heart width to coelom width of blue-fronted Amazon parrots (Amazona aestiva) with varying body condition scores. Thirty-five captive birds were included in the study and were allocated into one of three groups according to their respective body condition score: lean, ideal and obese. No differences were observed among the groups with regard to radiographic and tomographic measurements. Computed tomography enabled better identification of the structures of the cardiovascular system without interference from the overlying structures of the celomatic cavity observed in radiographic images. However, radiographic examinations should still be considered the standard screening method to identify cardiac alterations, such as increased or reduced organ dimensions. Standardizing the techniques and measurements performed in this study may serve as a basis for further research in the field.

Radiographic anatomy of the cockatiel (Nymphicus hollandicus) axial and appendicular skeleton.

Cockatiels are popular pets. Still, despite medical and surgical relevance, the radiographic anatomy of the cockatiel (Nymphicus hollandicus) skeleton, like that of different wild and exotic bird species, has seldom been described. This study set out to describe the radiographic anatomy of the cockatiel skeleton. Twelve adult male and nine adult female specimens were radiographed using a digital X-ray system and different views. The radiographic anatomy of these birds was similar to that of other Psittacidae. However, some particularities inherent to the target species were detected, such as the presence of four flexion zones in the skull (craniofacial, nasal, jugal arch and palatine), complete bony orbit comprising a suborbital arch, 34-38 vertebrae (10 or 11 cervical, 8 or 9 thoracic, 9 or 10 lumbosacral, 5 or 6 caudal vertebrae and a pygostyle comprising 2 fused vertebrae), eight or nine pairs of ribs and a notarium made up of fused T2-T6 vertebrae. Poor radiopacity of the notarium, ribs and respective uncinate processes, and synsacral vertebrae made demarcation of these structures difficult. The appendicular skeleton of the cockatiel was very similar to that of other Psittacidae, and there were no gender-related differences.

The endocast of the Night Parrot (Pezoporus occidentalis) reveals insights into its sensory ecology and the evolution of nocturnality in birds.

The Night Parrot (Pezoporus occidentalis) is a rare, nocturnal parrot species that has largely escaped scientific investigation due to its behaviour and habitat preferences. Recent field studies have revealed some insights into Night Parrot behaviour, but nothing is known of its sensory abilities. Here, we used µCT scans of an intact Night Parrot specimen to determine if its visual system shares similarities with other nocturnal species. The endocast of the Night Parrot revealed relatively small optic lobes and optic foramina, especially compared with closely related grass parakeets, but no apparent differences in orbit dimensions. Our data suggests that the Night Parrot likely has lower visual acuity than most other parrots, including its congener, the Eastern Ground Parrot (P. wallicus). We propose that the visual system of the Night Parrot might represent a compromise between the need to see under low light conditions and the visual acuity required to detect predators, forage, and fly. Based on the endocast and optic foramen measurements, the Night Parrot fits into a common pattern of decreased retinal input to the optic lobes in birds that should be explored more thoroughly in extant and extinct species.