Psittaciform birds exhibit novelties in jaw bone structure and musculature that are associated with strong bite forces. These features include an ossified arcus suborbitalis and the muscles ethmomandibularis and pseudomasseter. We analyse the jaw musculature of the monk parakeet (Myiopsitta monachus) to enable future studies aimed at understanding craniofacial development, morphology, function and evolution. We estimate bite force based on muscle dissections, physiological cross‐sectional area and skull biomechanical modelling. We also compare our results with available data for other birds and traced the evolutionary origin of the three novel diagnostic traits. Our results indicate that, in Myiopsitta, (i) the arcus suborbitalis is absent and the orbit is ventrally closed by an elongate processus orbitalis and a short ligamentum suborbitale; (ii) the ethmomandibularis muscle is a conspicuous muscle with two bellies, with its origin on the anterior portion of the septum interorbitale and insertion on the medial aspect of the mandible; (iii) the pseudomasseter muscle consists of some fibers arising from the m. adductor mandibulae externus superficialis, covering the lateral surface of the arcus jugalis and attaches by an aponeurotic sheet on the processus orbitalis; (iv) a well‐developed adductor mandibulae complex is present; (v) the bite force estimation relative to body mass is higher than that calculated for other non‐psittaciform species; and (vi) character evolution analysis revealed that the absence of the arcus suborbitalis and the presence of the m. pseudomassseter are the ancestral conditions, and mapping is inconclusive about presence of one or two bellies of the m. ethmomandibularis.
We describe the hindlimb myology of Milvago chimango. This member of the Falconidae: Polyborinae is a generalist and opportunist that can jump and run down prey on the ground, unlike Falconinae that hunt birds in flight and kill them by striking with its talons. Due to differences in the locomotion habits between the subfamilies, we hypothesized differences in their hindlimb myology. Gross dissections showed that the myology of M. chimango is concordant with that described of other falconids, except for the following differences: the m. flexor cruris medialis has one belly with a longitudinal division; the m. iliotibialis lateralis does not have a connection with the m. iliofibularis; the m. fibularis longus is strongly aponeurotic; the m. tibialis cranialis lacks an accessory tendons and the m. flexor hallucis longus has one place of origin, instead of two. The presence of the m. flexor cruris lateralis can be distinguished as it has been described absent for the Falconidae. We associated its presence with the predominant terrestrial habit of the M. chimango. Each muscle dissected was weighed and the relationship between flexors and extensors at each joint was assessed. The extensor muscles predominated in all joints in M. chimango. Among the flexors, the m. flexor hallucis longus was the heaviest, which could be related to the importance of the use of its talons to obtain food.
The ossification sequence of Myiopsitta monachus was determined. Myiopsitta has a similar sequence to other altricial birds, with delayed skeletons compared to precocial species. The hindlimbs ossify before the forelimbs, a condition that could be linked to altriciality. To determine the stability of the sequences of ossification across birds, we selected species of different groups of Aves and used event-pairing method and character mapping on a phylogeny. Our results show that the homogeneity in the development of birds was supported by 56.77% of the character states. Event-pair cracking phylogenetic method was applied to identify sequence heterochronies. Results reveal a high number of heterochronies and show that the long bones in limbs may behave as modules. In Myiopsitta, the ossa ectethmoidale and mesethmoidale ossify early. These bones provide the origin site of the Psittaciformes' novel adductor m. ethmomandibularis, associated with strong bite forces, and its acceleration in the sequence may correspond to the functional hypothesis. Also, the early appearance of some hyoid apparatus elements occurs, and could be related to the development of tongue in Psittaciformes and its role in handling food, and is in concordance with the functional and size hypothesis.
We studied the hindlimb myology of the monk parakeet (Myiopsitta monachus). Like all parrots, it has zygodactyl feet enabling perching, climbing, hanging, moving easily among trees, and handling food. Muscles were described and weighed, and physiological cross-sectional area (PCSA) of four flexors and one extensor was calculated. In comparison to other muscles, the M. tibialis cranialis and the M. fibularis brevis show increased development and high PCSA values, and therefore, large potential force production. Also, a large proportion of muscle mass was involved in flexing the digits. We hypothesize that these muscle traits are associated with the arboreal locomotion and food manipulation habits. In the monk parakeet, the M. extensor digitorum longus sends a branch to the hallux, and the connection between the M. flexor digitorum longus and the M. flexor hallucis longus is type I (Gadow's classification). We reaffirm the presence of the M. ambiens as a plesiomorphic condition that disappears in most members of the order. Among Psittaciformes, the M. fibularis brevis is stronger and the M. fibularis weaker in arboreal species than in basal terrestrial ones (e.g., Strigops).
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.
Knowledge about the embryonic stages of birds is important in answering many questions about development and evolution. We give the first description of 41 embryological stages of the monk parakeet (Myiopsitta monachus) on the basis of external morphology and comparison with the chicken. We also provide measurements of some external morphological characters (i.e. body mass, crown-rump, beak, forelimb, and third toe lengths) and perform comparisons with other precocial and altricial birds with the aim of identifying heterochronous developmental features. The following differences in the development of characters in the monk parakeet when compared with other birds were found: (1) delay of the feathers primordia, (2) wing buds initially greater than leg buds, (3) forelimbs and hindlimbs with similar relative size, (4) retroversion of the toe IV, (5) ventral curvature of the upper jaw, (6) positive regressions between stages and beak length with acceleration and higher values and III toe lengths with deceleration and lower values in the monk parakeet compared to the chicken. The growth pattern of the monk paraket Myiopsitta monachus could be influenced by some heterochronic processes like postdisplacement, acceleration and/or deceleration. Results of this research allow the standard identification of stages in different species of parrots, recognize similarities and differences between precocial (the chicken) and altricial species (Myiopsitta), and provide planning data for future studies.
The uropygial (preen) gland is a holocrine organ unique of Aves. Although several studies have been performed on the uropygial gland of different bird species, knowledge about this gland in Columbiformes is scarce. In order to fill this gap, we analysed in detail the external morphology and the histological and histochemical features of the uropygial gland of the Eared Dove (Zenaida auriculata) in a comparative context. The uropygial gland of the Eared Dove is characterized by its pear-like shape composed of two lobes, conical and naked papilla, tubule-alveolar adenomers, a large primary storage chamber (a feature also present in other terrestrial avian species), and reticular and elastic fibres in the capsule and connective tissue surrounding the adenomers. The histochemistry showed a positive reaction to periodic acid-Schiff, Alcian Blue 2.5 and several lectins, evidencing the presence of diverse glycoconjugates in this organ. Since the uropygial gland may be independently present or absent within Columbiformes, we also used character mapping on a molecular phylogeny to infer the character states of this gland at ancestral nodes to understand its evolutionary history. The analysis shows that the presence of the uropygial gland is the ancestral state for Columbiformes and that its loss occurred more than once independently. Keywords Preen gland • Zenaida auriculata • Histology • Histochemistry • Lectin • Ancestral state reconstruction ZusammenfassungDie Bürzeldrüse der Ohrflecktaube (Zenaida auriculata) und ihre evolutionsbiologische Geschichte innerhalb der Taubenvögel Die Bürzeldrüse ist eine nur bei Vögeln vorkommende holokrine Drüse. Obwohl es einige Untersuchungen dieser Drüse bei unterschiedlichen Vogelarten gibt, wissen wir nicht viel über sie bei Tauben. Um diese Lücke zu schließen, analysierten wir vergleichend und im Detail die äußerlichen morphologischen sowie die histologischen und histochemischen Eigenschaften der Bürzeldrüse von Ohrflecktauben (Zenaida auriculata). Ihre Bürzeldrüse ist durch ihre Birnenform charakterisiert und setzt sich zusammen aus zwei Lappen, einer konischen und nackten Papille, tubulealveolaren Adenomen, einer großen primären Speicherkapsel (die es auch bei anderen Landvögeln gibt), retikulären und elastischen Fasern in der Kapsel und Bindegewebe um die Adenomen herum. Die Histochemie zeigte positive Reaktionen auf PAS, Alcian-Blau 2,5 und mehrere Lektine, was auf das Vorhandensein von diversen Glykokonjugaten in diesem Organ hinweist. Da die Bürzeldrüse bei Tauben unabhängig voneinander vorhanden sein oder fehlen kann, betrachteten wir ihr Vorhandensein innerhalb des molekularen Stammbaums der Tauben, um die evolutionsbiologische Geschichte der Drüse zu verstehen. Die Analyse zeigt, dass das Vorhandensein der Bürzeldrüse bei Tauben entwicklungsgeschichtlich der Normalfall war und sie mehr als einmal unabhängig voneinander verlorenging.Communicated by L. Fusani.
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