A Comparison of Telencephalon Composition among Chickens, Junglefowl, and Wild Galliforms.

INTRODUCTION: Domestication is the process of modifying animals for human benefit through selective breeding in captivity. One of the traits that often diverges is the size of the brain and its constituent regions; almost all domesticated species have relatively smaller brains and brain regions than their wild ancestors. Although the effects of domestication on the brain have been investigated across a range of both mammal and bird species, almost nothing is known about the neuroanatomical effects of domestication on the world's most common bird: the chicken (Gallus gallus). METHODS: We compared the quantitative neuroanatomy of the telencephalon of white leghorn chickens with red junglefowl, their wild counterpart, and several wild galliform species. We focused specifically on the telencephalon because telencephalic regions typically exhibit the biggest differences in size in domesticate-wild comparisons. RESULTS: Relative telencephalon size was larger in chickens than in junglefowl and ruffed grouse (Bonasa umbellus). The relative size of telencephalic regions did not differ between chickens and junglefowl, but did differ in comparison with ruffed grouse. Ruffed grouse had larger hyperpallia and smaller entopallial, nidopallial, and striatal volumes than chickens and junglefowl. Multivariate analyses that included an additional three wild grouse species corroborated these findings: chicken and junglefowl have relatively larger nidopallial and striatal volumes than grouse. Conversely, the mesopallial and hyperpallial volumes tended to be relatively smaller in chickens and junglefowl. CONCLUSION: From this suite of comparisons, we conclude that chickens do not follow a pattern of widespread decreases in telencephalic region sizes that is often viewed as typical of domestication. Instead, chickens have undergone a mosaic of changes with some regions increasing and others decreasing in size, and there are few differences between chickens and junglefowl.

Symmetry breaking in the embryonic skin triggers directional and sequential plumage patterning.

The development of an organism involves the formation of patterns from initially homogeneous surfaces in a reproducible manner. Simulations of various theoretical models recapitulate final states of natural patterns, yet drawing testable hypotheses from those often remains difficult. Consequently, little is known about pattern-forming events. Here, we surveyed plumage patterns and their emergence in Galliformes, ratites, passerines, and penguins, together representing the three major taxa of the avian phylogeny, and built a unified model that not only reproduces final patterns but also intrinsically generates shared and varying directionality, sequence, and duration of patterning. We used in vivo and ex vivo experiments to test its parameter-based predictions. We showed that directional and sequential pattern progression depends on a species-specific prepattern: an initial break in surface symmetry launches a travelling front of sharply defined, oriented domains with self-organising capacity. This front propagates through the timely transfer of increased cell density mediated by cell proliferation, which controls overall patterning duration. These results show that universal mechanisms combining prepatterning and self-organisation govern the timely emergence of the plumage pattern in birds.

Allometric Scaling Rules of the Cerebellum in Galliform Birds.

Although the internal circuitry of the cerebellum is highly conserved across vertebrate species, the size and shape of the cerebellum varies considerably. Recent comparative studies have examined the allometric rules between cerebellar mass and number of neurons, but data are lacking on the numbers and sizes of Purkinje and granule cells or scaling of cerebellar foliation. Here, we investigate the allometric rules that govern variation in the volumes of the layers of the cerebellum, the numbers and sizes of Purkinje cells and granule cells and the degree of the cerebellar foliation across 7 species of galliform birds. We selected Galliformes because they vary greatly in body and brain sizes. Our results show that the molecular, granule and white matter layers all increase in volume at the same rate relative to total cerebellum volume. Both numbers and sizes of Purkinje cells increased with cerebellar volume, but numbers of Purkinje cells increased at a much faster rate than size. Granule cell numbers increased with cerebellar volume, but size did not. Sizes and numbers of Purkinje cells as well as numbers of granule cells were positively correlated with the degree of cerebellar foliation, but granule cell size decreased with higher degrees of foliation. The concerted changes among the volumes of cerebellar layers likely reflects the conserved neural circuitry of the cerebellum. Also, our data indicate that the scaling of cell sizes can vary markedly across neuronal populations, suggesting that evolutionary changes in cell sizes might be more complex than what is often assumed.

Selective attention in peacocks during assessment of rival males.

Males in many species compete intensely for access to females. In order to minimize costly interactions, they can assess their rivals' competitive abilities by evaluating traits and behaviors. We know little about how males selectively direct their attention to make these assessments. Using Indian peafowl (Pavo cristatus) as a model system, we examined how males visually assess their competitors by continuously tracking the gaze of freely moving peacocks during the mating season. When assessing rivals, peacocks selectively gazed toward the lower display regions of their rivals, including the lower eyespot and fishtail feathers, dense feathers, body and wings. Their attention was modified based on the rivals' behavior such that they spent more time looking at rivals when rivals were shaking their wings and moving. The results indicate that peacocks selectively allocate their attention during rival assessment. The gaze patterns of males assessing rivals were largely similar to those of females evaluating mates, suggesting that some male traits serve a dual function in both intra- and intersexual selection. However, males spent more time than females looking at the upper eyespots and this could indicate that the upper eyespots function more in close-up rival assessment than mate choice.

Morphometric traits of gizzard in relation to feeding habits of wild Sardinian partridges (Alectoris barbara barbara, Bonnaterre, 1790) with particular regard to clast selection.

Housing and feeding practices of wild birds for conservation management of biodiversity or restocking play a crucial role in determining the survival rates of animals when released into nature. Failure in coping with the environment might be one of the main flaws captive animals can experience when put into natural habitat. The present investigation aimed at exploring feeding habits and related morphometric traits of gizzard with respective content from wild partridges in comparison with captive ones. A total of 52 hunted wild Sardinian adult partridges (Alectoris barbara barbara Bonnaterre, 1790) were used. By comparison, 42 captive adult partridges reared in cages were enrolled. From each animal, the morphology of gizzard was investigated and respective content analysed for gross composition and taxonomical determination of fractions. Wet sieving analysis of each gizzard content was carried out (four-sieve towers with different mesh sizes: 1 mm, 500 µm, 250 µm and 125 µm), and relative and absolute weight of fresh filled and empty gizzards were recorded. Thickness of muscular layer of gizzard wall was measured by stereomicroscope. Carcass weight significantly (p < 0.05) differed between captive vs. wild partridges (478 ± 21 and 305 ± 35 g respectively). Post-mortem inspection highlighted gross morphological differences of gizzards between the two groups. Fresh weight of empty gizzards was 6.37 ± 0.80 vs. 11.25 ± 1.82 g, with average pH values of digesta 4.97 ± 0.11 vs. 4.38 ± 0.28 in captive vs. wild partridges respectively. Gizzard content from wild partridges accounted a 61.7% vs. 38.3% of biological vs. non-biological material proportions (DM basis). The non-biological material was mostly represented by lithic fragments and minerals (quartz, feldspar, calcite and mica) with specific peculiarities in terms of granulometry and morphometry. Feeding the captive partridges should point to support morphological and functional adaptation of gizzards to the feeding stuffs naturally available in the environment.

Testing the cranial evolutionary allometric 'rule' in Galliformes.

Recent comparative studies have indicated the existence of a common cranial evolutionary allometric (CREA) pattern in mammals and birds, in which smaller species have relatively smaller faces and bigger braincases than larger species. In these studies, cranial allometry was tested using a multivariate regression between shape (described using landmarks coordinates) and size (i.e. centroid size), after accounting for phylogenetic relatedness. Alternatively, cranial allometry can be determined by comparing the sizes of two anatomical parts using a bivariate regression analysis. In this analysis, a slope higher or lower than one indicates the existence of positive or negative allometry, respectively. Thus, in those species that support the CREA 'rule', positive allometry is expected for the association between face size and braincase size, which would indicate that larger species have disproportionally larger faces. In this study, I applied these two approaches to explore cranial allometry in 83 Galliformes (Aves, Galloanserae), ranging in mean body weight from 30 g to 2.5 kg. The multivariate regression between shape and centroid size revealed the existence of a significant allometric pattern resembling CREA, whereas the second analysis revealed a negative allometry for beak size and braincase size (i.e. contrary to the CREA 'rule', larger galliform species have disproportionally shorter beaks than smaller galliform species). This study suggests that the presence of CREA may be overestimated when using cranium size as the standard measurement.

Histological, histochemical and ultrastructural studies on Harderian and lacrimal glands of the Capercaillie (Tetrao urogallus major L.).

This study describes the macroscopic anatomy and the microscopic and ultrastructural features of the Harderian gland and lacrimal gland of the Capercaillies. It was conducted both on adult male and female Capercaillies. Tissue sections were stained with hematoxylin and eosin, azan trichrome, modified Mallory's trichrome, methyl green-pyronin Y, periodic acid-Schiff, alcian blue pH 2.5, aldehyde fuchsin and Hale's dialysed iron. The morphometric study of the Harderian and lacrimal glands indicated that they are both larger in male than in female Capercaillies. The histological analysis showed that the HG has a multilobar tubulo-alveolar structure with numerous lymphocytes and plasma cells. The LG has a multilobar tubulo-acinar structure without lymphocytes and plasma cells. The periodic acid-Schiff staining and alcian blue pH 2.5 staining demonstrated a mild positive reaction in the epithelial cells of the Harderian gland and weak positive reaction in the lacrimal gland. The HDI staining detected the presence of carboxylated acid mucopolysaccharides in the Harderian and lacrimal glands. Transmission electron microscopy revealed the presence of two types of secretory vesicles in the cytoplasm of both studied glands. It also showed that lipid droplets and glycogen granules were more abundant in the Harderian gland than in the lacrimal gland of this species.

Life-history differences in age-dependent expressions of multiple ornaments and behaviors in a lekking bird.

Age is a major factor explaining variation in life-history traits among individuals with typical patterns of increasing trait values early in life, maximum trait expression, and senescence. However, age-dependent variation in the expressions of sexually selected traits has received less attention, although such variation underpins differences in male competitive abilities and female preference, which are central to sexual selection. In contrast to previous studies focusing on single traits, we used repeated measures of seven sexually selected morphological and behavioral traits in male black grouse (Tetrao tetrix) to quantify the effects of age and life span on their expressions and quantified this variation in relation to male reproductive effort. Trait expression increased with age, but long-lived males had a slower increase and delayed maxima in trait values compared with short-lived males. There was evidence of terminal investment (increasing trait values during the last breeding season) in some traits and senescence in all traits. These trait dynamics were largely explained by the timing of male peak lekking effort. This study shows that fully understanding the variation in sexually selected traits and fitness benefits associated with sexual selection requires accounting for the complex interaction among individual age, life span, and the timing of individuals' investment in reproduction.

Guineafowl with a twist: asymmetric limb control in steady bipedal locomotion.

In avian bipeds performing steady locomotion, right and left limbs are typically assumed to act out of phase, but with little kinematic disparity. However, outwardly appearing steadiness may harbor previously unrecognized asymmetries. Here, we present marker-based XROMM data showing that guineafowl on a treadmill routinely yaw away from their direction of travel using asymmetrical limb kinematics. Variation is most strongly reflected at the hip joints, where patterns of femoral long-axis rotation closely correlate to degree of yaw divergence. As yaw deviations increase, hip long-axis rotation angles undergo larger excursions and shift from biphasic to monophasic patterns. At large yaw angles, the alternately striding limbs exhibit synchronous external and internal femoral rotations of substantial magnitude. Hip coordination patterns resembling those used during sidestep maneuvers allow birds to asymmetrically modulate their mediolateral limb trajectories and thereby advance using a range of body orientations.

Diet complexity in early life affects survival in released pheasants by altering foraging efficiency, food choice, handling skills and gut morphology.

Behavioural and physiological deficiencies are major reasons why reintroduction programmes suffer from high mortality when captive animals are used. Mitigation of these deficiencies is essential for successful reintroduction programmes. Our study manipulated early developmental diet to better replicate foraging behaviour in the wild. Over 2 years, we hand-reared 1800 pheasants (Phasianus colchicus), from 1 day old, for 7 weeks under different dietary conditions. In year one, 900 pheasants were divided into three groups and reared with (i) commercial chick crumb, (ii) crumb plus 1% live mealworm or (iii) crumb plus 5% mixed seed and fruit. In year two, a further 900 pheasants were divided into two groups and reared with (i) commercial chick crumb or (ii) crumb plus a combination of 1% mealworm and 5% mixed seed and fruit. In both years, the commercial chick crumb acted as a control treatment, whilst those with live prey and mixed seeds and fruits mimicking a more naturalistic diet. After 7 weeks reared on these diets, pheasants were released into the wild. Postrelease survival was improved with exposure to more naturalistic diets prior to release. We identified four mechanisms to explain this. Pheasants reared with more naturalistic diets (i) foraged for less time and had a higher likelihood of performing vigilance behaviours, (ii) were quicker at handling live prey items, (iii) were less reliant on supplementary feed which could be withdrawn and (iv) developed different gut morphologies. These mechanisms allowed the pheasants to (i) reduce the risk of predation by reducing exposure time whilst foraging and allowing more time to be vigilant; (ii) be better at handling and discriminating natural food items and not be solely reliant on supplementary feed; and (iii) have a better gut system to cope with the natural forage after the cessation of supplementary feeding in the spring. Learning food discrimination, preference and handling skills by the provision of a more naturalistic diet is essential prior to the release of pheasants in a reintroduction programme. Subsequent diet, foraging behaviour, gut morphology and digestive capabilities all work together as one nutritional complex. Simple manipulations during early development can influence these characteristics to better prepare an individual for survival upon release.

Sexing in guinea fowls (Numida meleagris).

Despite the potentials and contributions of guinea fowls to economic and social life in Ghana, accurate sex identification in these birds is still a major problem. Three hundred and sixty guinea fowls (180 birds per sex) were used in determining a more accurate and farmer friendly sexing technique. The sexing methods explored were vent, biometric, and molecular techniques. Vent sexing was accomplished by measuring phalli in 28 and 32-week-old birds, while biometric sexing involved the measurement of morphometric traits and data analyzed using discriminant function analysis. Molecular sexing was carried out by DNA extraction and subsequent PCR using the 2550F/2718R primer set. Females had a wider (P<0.05) pelvic inlet than male birds from first week of age until the end of the study, while the opposite was true for wattle length. However, wattle length differed (P<0.05) between both sexes after 4 weeks of age. Combining the biometric variables in a discriminant function, males could be distinguished from females with an accuracy of 94%. During molecular sexing, the P2/P8 primer set was not effective in sexing guinea fowls because it amplified a single band in both sexes and in the same manner. The sex of guinea fowls was properly determined using the primer set 2550F/2718R. Females produced 2 bands of 396 bp and 344 bp, while males only produced the larger band. Phallus size in the 2 sexes were distinguishable from 8 weeks of age, with males having longer and thicker (P<0.05) phalli than their female counterparts. Combining the 2 variables in a discriminate function, males and females could be distinguished with 98.3% accuracy. While the molecular method remains the most accurate sexing technique, the biometric method emerged as the most farmer friendly approach to sexing guinea fowls.

Long-axis rotation: a missing degree of freedom in avian bipedal locomotion.

Ground-dwelling birds are typically characterized as erect bipeds having hind limbs that operate parasagittally. Consequently, most previous research has emphasized flexion/extension angles and moments as calculated from a lateral perspective. Three-dimensional (3D) motion analyses have documented non-planar limb movements, but the skeletal kinematics underlying changes in foot orientation and transverse position remain unclear. In particular, long-axis rotation of the proximal limb segments is extremely difficult to measure with topical markers. Here, we present six degree of freedom skeletal kinematic data from maneuvering guineafowl acquired by marker-based XROMM (X-ray Reconstruction of Moving Morphology). Translations and rotations of the hips, knees, ankles and pelvis were derived from animated bone models using explicit joint coordinate systems. We distinguished sidesteps, sidestep yaws, crossover yaws, sidestep turns and crossover turns, but birds often performed a sequence of blended partial maneuvers. Long-axis rotation of the femur (up to 38 deg) modulated the foot's transverse position. Long-axis rotation of the tibiotarsus (up to 65 deg) also affected medio-lateral positioning, but primarily served to either re-orient a swing phase foot or yaw the body about a stance phase foot. Tarsometatarsal long-axis rotation was minimal, as was hip, knee and ankle abduction/adduction. Despite having superficially hinge-like joints, birds coordinate substantial long-axis rotations of the hips and knees to execute complex 3D maneuvers while striking a diversity of non-planar poses.

The effects of season and sex upon the morphology and material properties of keratin in the Svalbard rock ptarmigan (Lagopus muta hyperborea).

The material properties and morphologies of the modified integumentary organs of birds (the keratinous bills, claws and feathers) have evolved to withstand the variety of mechanical stresses imposed by their interaction with the environment. These stresses are likely to vary temporally in seasonal environments and may also differ between the sexes as a result of behavioural dimorphism. Here we investigate the morphology and material properties of the claws of male and female Svalbard ptarmigan (Lagopus muta hyperborea) during the summer and winter using nanoindentation. Despite differences in locomotor demands between the sexes and pronounced seasonal differences in environmental conditions, like ground substrate, ambient temperature and day length, there was no significant difference in Young׳s modulus or hardness between the seasons for each sex. However, when comparing males and females, female claws were significantly harder than those of males and both sexes had significantly wider claws during winter. We propose that wider claws may follow winter claw moulting as the claws are regrown and form an important part of the ptarmigan׳s snowshoe-like foot that is an adaptation to locomotion on snow. Future work focusing on growth rates and more broad measures of material properties in both captive and wild birds is required to determine the extent of seasonal and sex differences in the material properties of their keratinous structures.

Understanding sex differences in the cost of terrestrial locomotion.

Little is known regarding the physiological consequences of the behavioural and morphological differences that result from sexual selection in birds. Male and female Svalbard rock ptarmigans (Lagopus muta hyperborea) exhibit distinctive behavioural differences during the breeding season. In particular, males continuously compete for and defend territories in order to breed successfully, placing large demands on their locomotor system. Here, we demonstrate that male birds have improved locomotor performance compared with females, showing both a lower cost of locomotion (CoL) and a higher top speed. We propose that the observed sex differences in locomotor capability may be due to sexual selection for improved male performance. While the mechanisms underlying these energetic differences are unclear, future studies should be wary when pooling male and female data.

Environmental heterogeneity influences the reliability of secondary sexual traits as condition indicators.

Numerous studies have shown positive associations between ornaments and condition, as predicted by indicator models of sexual selection. However, this idea is continuously challenged by opposite results, which reveal our lack of full understanding of how sexual selection works. Environmental heterogeneity may explain such inconsistencies, but valid field tests of this idea are currently lacking. We first analysed the relationship between condition and ornament expression from nine populations over 7 years in a wild bird, the red grouse Lagopus lagopus scoticus. We then manipulated male aggressiveness at the population level by means of testosterone implants in a replicated field experiment. We found that the relationship between condition and ornamentation varied greatly between environments and became stronger when environmental conditions (ECs) were worse or when aggressiveness in the population was experimentally increased. Some ornaments may therefore reliably advertise a better condition only in adverse ECs. Considering environmental heterogeneity can help reconcile conflicting findings regarding the reliability of ornaments as indicators of condition and will help our understanding of sexual selection processes.

Interspecifc variation in eye shape and retinal topography in seven species of galliform bird (Aves: Galliformes: Phasianidae).

Eye morphology and the retinal topography of animals that live in either 'open' (e.g., grassland) or 'enclosed' (e.g., forest) terrestrial habitats show common adaptations to constraints imposed by these different habitat types. Although relationships between habitat and the visual system are well documented in most vertebrates, relatively few studies have examined this relationship in birds. Here, we compare eye shape and retinal topography across seven species from the family Phasianidae (Galliformes) that are diurnally active in either open or enclosed habitats. Species from enclosed habitats have significantly larger corneal diameters, relative to transverse diameters, than species from open habitats, which we predict serves to enhance visual sensitivity. Retinal topography, however, was similar across all seven species and consisted of a centrally positioned area centralis and a weak horizontal visual streak, with no discernible fovea. In the Japanese quail (Coturnix japonica), there was also a dorso-temporal extension of increased neuron density and, in some specimens, a putative area dorsalis. The total number of neurons in the retinal ganglion cell layer was correlated with retinal whole-mount area. Average and peak neuron densities were similar across species, with the exception of the Japanese quail, which had greater average and peak densities. Peak anatomical spatial resolving power was also similar among species, ranging from approximately 10-13 cycles/°. Overall, the pattern of retinal topography we found in phasianids is associated with ground-foraging in birds and presumably facilitates the identification of small food items on the ground as well as other visually guided behaviors, irrespective of habitat type.

Anatomical and histological structure of the tongue and histochemical characteristics of the lingual salivary glands in the Chukar partridge (Alectoris chukar, Gray 1830).

1. The aim of the study was to examine the morphology of the tongue and the histochemical features of the lingual salivary glands in this species. 2. The tongue was elongated, terminating in a rather sharp, dagger-like apex. On the surface of the tongue and situated between the body and root of the tongue, two rows of conical papillae, the sharp apices of which pointed towards the posterior part of the tongue, were observed. The keratinised epithelium lining the dorsal surface lacked typical gustatory papillae. However, it was observed that taste buds were present in the epithelium of the lingual body and root. The tongue was supported by a structure composed of hyaline cartilage, the paraglossum, which extended from the lingual root to the apex. Simple branched tubular glands, which were encapsulated by connective tissue, were embedded within the submucosa in the body (anterior salivary glands) and root (posterior salivary glands) of the tongue. It was observed that the secretion of the lingual glands contained neutral mucins, proteoglycans containing carboxylic acid, weak and strong sulphated groups, N-acetylated sialomucins, but lacked glycogen. 3. It was demonstrated that, the general morphological features, papillary distribution of the tongue and the histological structure of the mucosa epithelium and the supportive elements displayed similarity to those of other domestic avian species. It was also determined that, in view of the particular feeding types, in the partridge, the presence of the papillary crest was not correlated with diet.

Body weight and some biometrical traits of ring-necked pheasants (Phasianus colchicus) at different ages.

Although the growth of pheasants is an important issue, there are few publications providing information on body weight and morphometrics of pheasants at different ages. The aim of this study was to investigate body weight, tarsus length, wing length and wingspan in farmed pheasants aged 3-16 weeks. The study was carried out on 50 pheasants (31 females and 19 males). All one-day old birds were marked and kept until the 16th week of life. In the 3rd 8 h 12th and 16th weeks of life all pheasants were weighted and tarsus length, wing length and wingspan were measured. Male chicks appeared to be significantly heavier than females (P< or = 0.01) already in the third week of life. Also their wings were longer and their wingspan attained higher values (P < or = 0.05). Absolute gains of the majority of traits also differed between sexes (being larger in males) in all studied periods. Values of body weight, tarsus length and wingspan noted in the third week of life were significantly correlated with all later measurements. Thus body weight and some biometrical traits of pheasant chicks measured already at the 3rd week of life could be applied as predictors of future body weight and size.

The ornament-condition relationship varies with parasite abundance at population level in a female bird.

Environmental heterogeneity is expected to create variation in the ornament-condition relationship. This topic has been studied in males with less attention being given to females. Here, we explore inter-population variation in the relationship between the size of a male-like trait, supra-orbital combs, and body mass in female red grouse Lagopus lagopus scoticus. We used the abundance of the nematode Trichostrongylus tenuis, a parasite with strong negative effects on this species, as a proxy of environmental conditions. We studied six populations over 5 years and showed that the comb size-body mass relationship varied with mean parasite abundance, with stronger ornament-condition relationships in populations with higher parasite infection levels. Our study supports the idea that environmental conditions, and in particular parasite infection levels, may affect the reliability of female ornaments as condition indicators.

Morphology of the tongue and laryngeal entrance in the Common pheasant (Phasianus colchicus); a light and scanning electron microscopic study.

The present work was carried out to describe morphological features of the tongue and laryngeal entrance of the Common pheasantPhasianus colchicus, using light and scanning electron microscopy. The lingual apex was taper-shaped with a shallow median sulcus. A V-shaped papillary crest with 15-20 caudally directed conical papillae was located between the radix and corpus linguae. The second row of papillae consisting of three caudolaterally directed conical papillae was arranged caudal to the main papillary crest. The mons laryngealis was equipped caudally with a double-layer of large conical papillae as pharyngeal crest, and few small conical papillae were seen around the glottis. Scanning electron microscope (SEM) observation showed that the apex linguae and corpus were covered with compact filiform papillae, while the radix linguae were free of papillae. Numerous openings of salivary gland can be seen on the radix linguae and mons laryngealis at higher magnification. On histological sections, all parts of the tongue and mons laryngealis were covered with a keratinised stratified squamous epithelium. The keratinised layer became thinner rostro-caudally. The simple alveolar salivary glands were distributed in the submucosa from the apex linguae to the radix linguae as well as the mons laryngealis. PAS staining revealed mucin-secreting activity of this glands. According to the findings, although the tongue and laryngeal entrance of the Common pheasant is characteristics of a galliform bird, it has some features similar to passeriformes and falconiformes.