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1

Botelho, João Francisco, Daniel Smith-Paredes, Daniel Nuñez-Leon, Sergio Soto-Acuña, and Alexander O. Vargas. "The developmental origin of zygodactyl feet and its possible loss in the evolution of Passeriformes." Proceedings of the Royal Society B: Biological Sciences 281, no. 1788 (2014): 20140765. http://dx.doi.org/10.1098/rspb.2014.0765.

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The zygodactyl orientation of toes (digits II and III pointing forwards, digits I and IV pointing backwards) evolved independently in different extant bird taxa. To understand the origin of this trait in modern birds, we investigated the development of the zygodactyl foot of the budgerigar (Psittaciformes). We compared its muscular development with that of the anisodactyl quail (Galliformes) and show that while the musculus abductor digiti IV (ABDIV) becomes strongly developed at HH36 in both species, the musculus extensor brevis digiti IV (EBDIV) degenerates and almost disappears only in the budgerigar. The asymmetric action of those muscles early in the development of the budgerigar foot causes retroversion of digit IV (dIV). Paralysed budgerigar embryos do not revert dIV and are anisodactyl. Both molecular phylogenetic analysis and palaeontological information suggest that the ancestor of passerines could have been zygodactyl. We followed the development of the zebra finch (Passeriformes) foot muscles and found that in this species, both the primordia of the ABDIV and of the EBDIV fail to develop. These data suggest that loss of asymmetric forces of muscular activity exerted on dIV, caused by the absence of the ABDIV, could have resulted in secondary anisodactyly in Passeriformes.
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2

Mayr, Gerald. "A new family of Eocene zygodactyl birds." Senckenbergiana lethaea 78, no. 1-2 (1998): 199–209. http://dx.doi.org/10.1007/bf03042769.

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3

Lai, Anna Chiara, and Paola Loreti. "Self-similar control systems and applications to zygodactyl bird's foot." Networks and Heterogeneous Media 10, no. 2 (2015): 401–19. http://dx.doi.org/10.3934/nhm.2015.10.401.

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4

Senthil, Pavan, Om Vishanagra, John Sparkman, Peter Smith, and Albert Manero. "Design and Assessment of Bird-Inspired 3D-Printed Models to Evaluate Grasp Mechanics." Biomimetics 9, no. 4 (2024): 195. http://dx.doi.org/10.3390/biomimetics9040195.

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Adapting grasp-specialized biomechanical structures into current research with 3D-printed prostheses may improve robotic dexterity in grasping a wider variety of objects. Claw variations across various bird species lend biomechanical advantages for grasping motions related to perching, climbing, and hunting. Designs inspired by bird claws provide improvements beyond a human-inspired structure for specific grasping applications to offer a solution for mitigating a cause of the high rejection rate for upper-limb prostheses. This research focuses on the design and manufacturing of two robotic test devices with different toe arrangements. The first, anisodactyl (three toes at the front, one at the back), is commonly found in birds of prey such as falcons and hawks. The second, zygodactyl (two toes at the front, two at the back), is commonly found in climbing birds such as woodpeckers and parrots. The evaluation methods for these models included a qualitative variable-object grasp assessment. The results highlighted design features that suggest an improved grasp: a small and central palm, curved distal digit components, and a symmetrical digit arrangement. A quantitative grip force test demonstrated that the single digit, the anisodactyl claw, and the zygodactyl claw designs support loads up to 64.3 N, 86.1 N, and 74.1 N, respectively. These loads exceed the minimum mechanical load capabilities for prosthetic devices. The developed designs offer insights into how biomimicry can be harnessed to optimize the grasping functionality of upper-limb prostheses.
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Lockley, Martin G., Rihui Li, Jerald D. Harris, Masaki Matsukawa, and Mingwei Liu. "Earliest zygodactyl bird feet: evidence from Early Cretaceous roadrunner-like tracks." Naturwissenschaften 94, no. 8 (2007): 657–65. http://dx.doi.org/10.1007/s00114-007-0239-x.

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6

Necas, Petr. "Nečas, P. (2020) Chameleodactyly: New term to describe the unique arrangement of digits in chameleons (Reptilia: Chamaeleonidae). – Archaius 1 (1): 4 – 5." Archaius 1, no. 1 (2020): 4–5. https://doi.org/10.5281/zenodo.3751185.

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7

Carril, Julieta, Claudio G. Barbeito, and Claudia P. Tambussi. "Making a parrot zygodactyl foot: Osteology and morphogenesis of the tarsometatarsus in the monk parakeet (Myiopsitta monachus)." Zoology 144 (February 2021): 125877. http://dx.doi.org/10.1016/j.zool.2020.125877.

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8

Mayr, Gerald. "A reassessment of Eocene parrotlike fossils indicates a previously undetected radiation of zygodactyl stem group representatives of passerines (Passeriformes)." Zoologica Scripta 44, no. 6 (2015): 587–602. http://dx.doi.org/10.1111/zsc.12128.

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9

Mayr, Gerald. "A partial skeleton of a new species of Tynskya Mayr, 2000 (Aves, Messelasturidae) from the London Clay highlights the osteological distinctness of a poorly known early Eocene “owl/parrot mosaic”." PalZ 95, no. 2 (2021): 337–57. http://dx.doi.org/10.1007/s12542-020-00541-8.

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AbstractTynskya eocaena is an early Eocene bird with a raptor-like skull and semi-zygodactyl feet, whose description is based on a skeleton from the North American Green River Formation. In the present study, three-dimensionally preserved bones of a new species of Tynskya, T. waltonensis, are reported from the London Clay of Walton-on-the-Naze (Essex, UK). The fossils belong to a single individual and provide new insights into the skeletal morphology of messelasturids. In particular, they reveal unusual vertebral specializations, with the cervical vertebrae having concave rather than saddle-shaped caudal articulation facets and the caudalmost thoracic vertebra being platycoelous (flat articular surfaces). The very deep mandible and a derived morphology of the ungual phalanges support a sister group relationship between Tynskya and the taxon Messelastur (Messelasturidae). Phylogenetic analyses of an emended data matrix did not conclusively resolve the higher-level affinities of messelasturids and the closely related halcyornithids, with both taxa sharing derived characters with only distantly related extant taxa (Accipitriformes, Strigiformes, Falconiformes, and Psittaciformes). An analysis that was constrained to a molecular scaffold, however, recovered messelasturids as the sister taxon of a clade including psittaciform and passeriform birds. The derived morphologies of the mandible and cervical vertebrae suggest specialized feeding adaptations of Tynskya, and messelasturids may have exploited a feeding niche, which is no longer available to extant birds.
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10

J, Violet Beaulah, P. Sridevi, K. S. Ravali, P. Dharani, S. Rajathi, and T. A. Kannan. "Hindlimb Skeletal Structure of the Green-winged Macaw: An Anatomical Study." UTTAR PRADESH JOURNAL OF ZOOLOGY 45, no. 10 (2024): 26–33. http://dx.doi.org/10.56557/upjoz/2024/v45i104042.

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In birds, the forelimb bones undergo modification to facilitate muscle attachment for flight, enabling the movement of wings up and down. Conversely, hindlimb bones primarily support walking and perching functions, necessitating evolutionary adaptations in their structure. Therefore, this study aims to document the gross anatomical features of hind limb bones in Green-winged Macaws, including the femur, tibio-tarsus, tarsometatarsus, and digits. The bones were sourced from six Green-winged Macaw carcasses undergoing post-mortem examination at the Department of Veterinary Pathology, Madras Veterinary College, Chennai. Preparation was conducted using the wet maceration technique. In the femur, the proximal end displayed a large, well-defined spherical head medially, accompanied by a small depression called the fovea capitis, and a distinct neck. The tibio-tarsus exhibited a small roughened area on its lateral border for fibula attachment, with a larger medial and smaller lateral condyle at the proximal extremity, along with a linear cinemal crest along the medial border. The fibula's distal extremity tapered into a free point, articulating at the caudolateral aspect of the tarsometatarsus. The tarsometatarsus displayed fused distal tarsals with metatarsals II, III, and IV, while Metatarsus I remained a separate bone, forming the base of the first toe. The proximal extremity of the tarsometatarsus featured two large concave articular facets for condyles from the distal extremity of the tibio-tarsus, and a facet for the distal end of the fibula caudolaterally. The Macaw's hind limb consisted of four digits, forming a zygodactyl foot arrangement.
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11

Mayr, Gerald. "A remarkably complete skeleton from the London Clay provides insights into the morphology and diversity of early Eocene zygodactyl near-passerine birds." Journal of Systematic Palaeontology 18, no. 22 (2020): 1891–906. http://dx.doi.org/10.1080/14772019.2020.1862930.

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12

Sustaita, Diego, Yuri Gloumakov, Leah R. Tsang, and Aaron M. Dollar. "Behavioral correlates of semi-zygodactyly in Ospreys (Pandion haliaetus) based on analysis of internet images." PeerJ 7 (February 5, 2019): e6243. http://dx.doi.org/10.7717/peerj.6243.

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Ospreys are renowned for their fishing abilities, which have largely been attributed to their specialized talon morphology and semi-zygodactyly−the ability to rotate the fourth toe to accompany the first toe in opposition of toes II and III. Anecdotal observations indicate that zygodactyly in Ospreys is associated with prey capture, although to our knowledge this has not been rigorously tested. As a first pass toward understanding the functional significance of semi-zygodactyly in Ospreys, we scoured the internet for images of Osprey feet in a variety of circumstances. From these we cross-tabulated the number of times each of three toe configurations (anisodactylous, zygodactylous, and an intermediate condition between these) was associated with different grasping scenarios (e.g., grasping prey or perched), contact conditions (e.g., fish, other objects, or substrate), object sizes (relative to foot size), and grasping behaviors (e.g., using one or both feet). Our analysis confirms an association between zygodactyly and grasping behavior; the odds that an osprey exhibited zygodactyly while grasping objects in flight were 5.7 times greater than whilst perched. Furthermore, the odds of zygodactyly during single-foot grasps were 4.1 times greater when pictured grasping fish compared to other objects. These results suggest a functional association between predatory behavior and zygodactyly and has implications for the selective role of predatory performance in the evolution of zygodactyly more generally.
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13

Niblock, Aaron, Dominic Og McConville, and Patrick John Morrison. "Zygodactyly is strongly associated with Acute Myeloid Leukaemia." British Journal of Haematology 177, no. 4 (2016): 659–60. http://dx.doi.org/10.1111/bjh.14096.

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14

Mayr, Gerald, and Nikita Zelenkov. "New specimens of zygodactylid birds from the middle Eocene of Messel, with description of a new species of Primozygodactylus." Acta Palaeontologica Polonica 54, no. 1 (2009): 15–20. https://doi.org/10.4202/app.2009.B103.

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Mayr, Gerald, Zelenkov, Nikita (2009): New specimens of zygodactylid birds from the middle Eocene of Messel, with description of a new species of Primozygodactylus. Acta Palaeontologica Polonica 54 (1): 15-20, DOI: 10.4202/app.2009.B103, URL: http://www.bioone.org/doi/abs/10.4202/app.2009.B103
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15

McConville, Dominic O., G. Pooler Archbold, Anthony Lewis, and Patrick J. Morrison. "Zygodactyly (Syndactyly Type A1) Associated With Midfoot Charcot Neuropathy and Diabetes." Diabetes Care 41, no. 5 (2018): e74-e75. http://dx.doi.org/10.2337/dc18-0011.

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16

Smith, N. Adam, Aj M. DeBee, and Julia A. Clarke. "Systematics and phylogeny of the Zygodactylidae (Aves, Neognathae) with description of a new species from the early Eocene of Wyoming, USA." PeerJ 6 (June 25, 2018): e4950. http://dx.doi.org/10.7717/peerj.4950.

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Zygodactylidae are an extinct lineage of perching birds characterized by distinct morphologies of the foot and wing elements. Although the clade has a complex taxonomic history, current hypotheses place Zygodactylidae as the sister taxon to Passeriformes (i.e., songbirds). Given the rather sparse fossil record of early passeriforms, the description of zygodactylid taxa is important for inferring potentially ancestral states in the largest radiation of living birds (i.e., the ∼6,000 species of extant passeriforms). Despite the exceptional preservation of many specimens and considerable species diversity in Zygodactylidae, the relationships among species have not been previously evaluated in a phylogenetic context. Herein, we review the fossil record of Zygodactylidae from North America and describe five new well-preserved fossils from the early Eocene Green River Formation of Wyoming. Two specimens are identified as representing a new species and the first records of the taxonZygodactylusoutside Europe. Anatomical comparisons with previously named taxa and the results of phylogenetic analysis including newly described specimens and previously named zygodactylid taxa provide the first hypothesis of the species-level relationships among zygodactylids. The monophyly of Zygodactylidae is supported in these new analyses. However, the monophyly ofPrimozygodactylusand the taxonomic distinction betweenZygodactylusandEozygodactylusremain unresolved and would likely benefit from the description of additional specimens.
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17

Ksepka, Daniel T., Thomas A. Stidham, and Thomas E. Williamson. "Early Paleocene landbird supports rapid phylogenetic and morphological diversification of crown birds after the K–Pg mass extinction." Proceedings of the National Academy of Sciences 114, no. 30 (2017): 8047–52. http://dx.doi.org/10.1073/pnas.1700188114.

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Evidence is accumulating for a rapid diversification of birds following the K–Pg extinction. Recent molecular divergence dating studies suggest that birds radiated explosively during the first few million years of the Paleocene; however, fossils from this interval remain poorly represented, hindering our understanding of morphological and ecological specialization in early neoavian birds. Here we report a small fossil bird from the Nacimiento Formation of New Mexico, constrained to 62.221–62.517 Ma. This partial skeleton represents the oldest arboreal crown group bird known. Phylogenetic analyses recoveredTsidiiyazhi abinigen. et sp. nov. as a member of the Sandcoleidae, an extinct basal clade of stem mousebirds (Coliiformes). The discovery ofTsidiiyazhipushes the minimum divergence ages of as many as nine additional major neoavian lineages into the earliest Paleocene, compressing the duration of the proposed explosive post–K–Pg radiation of modern birds into a very narrow temporal window parallel to that suggested for placental mammals. Simultaneously,Tsidiiyazhiprovides evidence for the rapid morphological (and likely ecological) diversification of crown birds. Features of the foot indicate semizygodactyly (the ability to facultatively reverse the fourth pedal digit), and the arcuate arrangement of the pedal trochleae bears a striking resemblance to the conformation in owls (Strigiformes). Inclusion of fossil taxa and branch length estimates impacts ancestral state reconstructions, revealing support for the independent evolution of semizygodactyly in Coliiformes, Leptosomiformes, and Strigiformes, none of which is closely related to extant clades exhibiting full zygodactyly.
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18

Mayr, Gerald, and Nikita Zelenkov. "New Specimens of Zygodactylid Birds from the Middle Eocene of Messel, with Description of a New Species ofPrimozygodactylus." Acta Palaeontologica Polonica 54, no. 1 (2009): 15–20. http://dx.doi.org/10.4202/app.2009.b103.

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19

Mumtaz, Sara, Esra Yıldız, Karmoon Lal, Aslıhan Tolun, and Sajid Malik. "Complex postaxial polydactyly types A and B with camptodactyly, hypoplastic third toe, zygodactyly and other digit anomalies caused by a novel GLI3 mutation." European Journal of Medical Genetics 60, no. 5 (2017): 268–74. http://dx.doi.org/10.1016/j.ejmg.2017.03.004.

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20

Mayr, Gerald, S. Bruce Archibald, Gary W. Kaiser, and Rolf W. Mathewes. "Early Eocene (Ypresian) birds from the Okanagan Highlands, British Columbia (Canada) and Washington State (USA)." Canadian Journal of Earth Sciences 56, no. 8 (2019): 803–13. http://dx.doi.org/10.1139/cjes-2018-0267.

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We survey the known avian fossils from Ypresian (early Eocene) fossil sites of the North American Okanagan Highlands, mainly in British Columbia (Canada). All specimens represent taxa that were previously unknown from the Eocene of far-western North America. Wings from the McAbee site are tentatively referred to the Gaviiformes and would constitute the earliest fossil record of this group of birds. A postcranial skeleton from Driftwood Canyon is tentatively assigned to the Songziidae, a taxon originally established for fossils from the Ypresian of China. Two skeletons from Driftwood Canyon and the McAbee site are tentatively referred to Coliiformes and Zygodactylidae, respectively, whereas three further fossils from McAbee, Blakeburn, and Republic (Washington, USA) are too poorly preserved for even a tentative assignment. The specimens from the Okanagan Highlands inhabited relatively high paleoaltitudes with microthermal climates (except Quilchena: lower mesothermal) and mild winters, whereas most other Ypresian fossil birds are from much warmer lowland paleoenvironments with upper mesothermal to megathermal climates. The putative occurrence of a gaviiform bird is particularly noteworthy because diving birds are unknown from other lacustrine Ypresian fossil sites of the Northern Hemisphere. The bones of the putative zygodactylid show a sulphurous colouration, and we hypothesize that this highly unusual preservation may be due to the metabolic activity of sulphide-oxidizing bacteria.
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21

Mayr, Gerald. "New species of Primozygodactylus from Messel and the ecomorphology and evolutionary significance of early Eocene zygodactylid birds (Aves, Zygodactylidae)." Historical Biology 29, no. 7 (2016): 875–84. http://dx.doi.org/10.1080/08912963.2016.1261135.

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22

Lynch, John D. "THE RELATIONSHIPS OF AN ENSEMBLE OF GUATEMALAN AND MEXICAN FROGS (ELEUTHERODACTYLUS: LEPTODACTYLIDAE: AMPHIBIA)." Revista de la Academia Colombiana de Ciencias Exactas, Físicas y Naturales 24, no. 90 (2024): 129–56. http://dx.doi.org/10.18257/raccefyn.24(90).2000.2843.

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Eleutherodactylus daryi and E. greggi are shown to be species of the Eleutherodactylus gollmeri group which is defined on the basis of having a synapomorphy (fusion of sacrum and presacral vertebrae). Eleutherodactylus omiltemanus is a species of the rhodopis Series rather than thefitzingeri Series. The Eleutherodactylus omiltemanus group disappears because ali of its contained members are placed in other species groups. Eleutherodactylus saltator is a synonym of E. mexicanus. Eleutherodactylus rhodopis sensu lato contains two species, a larger upland E. rhodopis and the more widely distibuted lowland E. loki. The rhodopis Series, containing the rhodopis group, is not definable because no synapomorphies are apparent. Eleutherodactylus occidentalis and E. omiltemanus are assigned to this "group. " Within this "group " there is a cluster of species having black mesorchia (E. hobartsmithi, E. mexicanus, E. pygmaeus, and E. sarton), one having a shortened inner toe (E. jota and E. podiciferus), and one having conical subarticular tubercles (E. bransfordii, E. lauraster, E. loki, E. occidentalis, E. omiltemanus, E. rhodopis, and E. stejnegerianus). Sixty of the 84 essentially Mesoamerican species of the subgenus Craugastor are placed in a monophyletic subunit based on the common presence of marked sexual dimorphism in tympanum size. The subunit includes the nine species ofthe gollmeri group, the thirteen species of the rhodopis group, and 38 additional species (the three species of the biporcatus group, the eleven species of the milesi group, eight species of thefitzingeri group, 15 species of the rugulosus group, and E. uno). The 24 species of Craugastor not included in that clade are the 13 species of the alfredi group (including E. bocourti), the two species of the augusti group, the two species of the bufoniformis group, three species removed from thefitzingeri group (E. andi, E. cuaquero, and E. emcelae), and four species removed from the rugulosus group (E. anatipes, E. anomalus, E. cheiroplethus, and E. zygodactylus).
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23

Anderson, C. Jane, Eric A. Tillman, William P. Bukoski, et al. "A novel parakeet‐selective feeder for control of invasive psittacines." Wildlife Society Bulletin, September 3, 2023. http://dx.doi.org/10.1002/wsb.1483.

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AbstractOver 40 species of parrots, members of order Psittaciformes, have established nonnative populations globally. Monk parakeets (Myiopsitta monachus) are among the most invasive bird species worldwide. In their introduced range, populations of monk parakeets have caused negative impacts on native species, habitats, economies, and human safety. Lethal population management has been complicated by the intelligence of monk parakeets, as they quickly alter behavior to avoid risks. Further, lethal control programs have been halted due to public controversy, as parakeets are highly charismatic. The contraceptive DiazaCon has been demonstrated to effectively reduce fertility in monk parakeets and other psittacines. In field applications, chemical control agents (e.g., toxicants and contraceptives) must be delivered in a manner that prohibits access by nontarget species. We developed and tested a parakeet‐selective feeder. The feeder allows access by parakeets and limits access by nontarget bird species by lowering a wire exclusion curtain around the feeder, requiring a zygodactyl toe arrangement to access food. We tested the parakeet‐selective feeder in trials with captive and free‐ranging monk parakeets and nontarget species in Florida, USA. Monk parakeets successfully accessed food from the parakeet‐selective feeder throughout the study. The mean number of daily feeder uses by nontarget species decreased from a high of nearly 16 uses per day when the exclusion curtain was not implemented to <1 use per day when implemented. Our findings suggest the parakeet‐selective feeder is a promising tool for delivery of bait treated with chemical control agents to manage monk parakeets and other nonnative parakeet populations, but implementation success will likely vary by target species, location, local faunal diversity, and availability of alternative forage.
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Lúcio de Souza, Mayara Oliveira, Fernanda Menezes de Lima, Liliane da Silva Barbosa, Maria Eduarda Brêda Accorsi, Mariana Lima dos Santos, and Müller Ribeiro-Andrade. "Proposal for the use of morphometry as a sexing method for psittacine species Amazona amazonica, Amazona aestiva , Ara chloropterus , Anodorhynchus hyacinthinus , and Ara ararauna." Avian Biology Research, April 28, 2025. https://doi.org/10.1177/17581559251336691.

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Psittacides are birds characterized by a curved beak, zygodactyl feet and colorful feathers, with species varying in size and without apparent sexual dimorphism. They are among the birds most threatened with extinction, with around 31% of species in the Neotropical region in this condition. Reproduction of these birds in captivity is crucial for the maintenance and recovery of populations, with sex identification being essential in this process. Given the difficulty of differentiating between males and females, it is proposed to use external morphometry to determine sex in a population of parrots, correlating it with the measurement of five parameters: length of the culmen, tomium, depth of the beak, length of the head and cloaca opening. Sex was determined by DNA testing, where females presented two copies of genes amplified by PCR (one from each chromosome, W and Z) and males presented only one copy (Z chromosome). 40 parrot specimens of different species were used: 7 Amazona amazonica , 8 Amazona aestiva , 9 Ara chloropterus , 5 Anodorhynchus hyacinthinus , 11 Ara ararauna . Of the total, 55% (22/40) were males and 45% (18/40) were females. Among the parameters evaluated, there were no significant variances between the sexes of the species studied, with the exception of one variable vent opening of one species Ara ararauna . Additionally, the lack of information about the age range of the animals meant that some individuals showed similarities, as in the case of A. amazonica , possibly linked to the stage of development. Thus, in addition to verifying that not all species can be subjected to the morphometric technique, the development phase also interferes with its use. It is concluded that among the species studied, only Ara ararauna is possible to determine the sex by measuring the cloaca opening, which showed significant differences, with no divergence from other biometric parameters.
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Hieronymus, Tobin L., David A. Waugh, and Julia A. Clarke. "A new zygodactylid species indicates the persistence of stem passerines into the early Oligocene in North America." BMC Evolutionary Biology 19, no. 1 (2019). http://dx.doi.org/10.1186/s12862-018-1319-6.

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