Pygostyles
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Pygostyle is a skeletal condition in which the final few
caudal Caudal may refer to: Anatomy * Caudal (anatomical term) (from Latin ''cauda''; tail), used to describe how close something is to the trailing end of an organism * Caudal artery, the portion of the dorsal aorta of a vertebrate that passes into th ...
vertebra Each vertebra (: vertebrae) is an irregular bone with a complex structure composed of bone and some hyaline cartilage, that make up the vertebral column or spine, of vertebrates. The proportions of the vertebrae differ according to their spina ...
e are fused into a single
ossification Ossification (also called osteogenesis or bone mineralization) in bone remodeling is the process of laying down new bone material by cells named osteoblasts. It is synonymous with bone tissue formation. There are two processes resulting in t ...
, supporting the tail
feather Feathers are epidermal growths that form a distinctive outer covering, or plumage, on both avian (bird) and some non-avian dinosaurs and other archosaurs. They are the most complex integumentary structures found in vertebrates and an exa ...
s and musculature. In modern
bird Birds are a group of warm-blooded vertebrates constituting the class (biology), class Aves (), characterised by feathers, toothless beaked jaws, the Oviparity, laying of Eggshell, hard-shelled eggs, a high Metabolism, metabolic rate, a fou ...
s, the
rectrices Flight feathers (''Pennae volatus'') are the long, stiff, asymmetrically shaped, but symmetrically paired pennaceous feathers on the Bird wing, wings or tail of a bird; those on the wings are called remiges (), singular remex (), while those ...
attach to these. The pygostyle is the main component of the uropygium, a structure colloquially known as the bishop's nose, parson's nose, pope's nose, or sultan's nose. This is the fleshy protuberance visible at the posterior end of a
bird Birds are a group of warm-blooded vertebrates constituting the class (biology), class Aves (), characterised by feathers, toothless beaked jaws, the Oviparity, laying of Eggshell, hard-shelled eggs, a high Metabolism, metabolic rate, a fou ...
(most commonly a
chicken The chicken (''Gallus gallus domesticus'') is a domesticated subspecies of the red junglefowl (''Gallus gallus''), originally native to Southeast Asia. It was first domesticated around 8,000 years ago and is now one of the most common and w ...
or
turkey Turkey, officially the Republic of Türkiye, is a country mainly located in Anatolia in West Asia, with a relatively small part called East Thrace in Southeast Europe. It borders the Black Sea to the north; Georgia (country), Georgia, Armen ...
) that has been dressed for
cooking Cooking, also known as cookery or professionally as the culinary arts, is the art, science and craft of using heat to make food more palatable, digestible, nutritious, or Food safety, safe. Cooking techniques and ingredients vary widely, from ...
. It has a swollen appearance because it also contains the
uropygial gland The uropygial gland, informally known as the preen gland or the oil gland, is a bilobed sebaceous gland possessed by the majority of birds used to distribute the gland's oil through the plumage by means of Preening (bird), preening. It is locate ...
that produces
preen oil The uropygial gland, informally known as the preen gland or the oil gland, is a bilobed sebaceous gland possessed by the majority of birds used to distribute the gland's oil through the plumage by means of preening. It is located dorsally at th ...
.


Evolution

Pygostyles probably began to evolve very early in the
Cretaceous The Cretaceous ( ) is a geological period that lasted from about 143.1 to 66 mya (unit), million years ago (Mya). It is the third and final period of the Mesozoic Era (geology), Era, as well as the longest. At around 77.1 million years, it is the ...
period, perhaps 140 – 130 million years ago. The earliest known
species A species () is often defined as the largest group of organisms in which any two individuals of the appropriate sexes or mating types can produce fertile offspring, typically by sexual reproduction. It is the basic unit of Taxonomy (biology), ...
to have evolved a pygostyle were members of the
Confuciusornithidae Confuciusornithidae is an extinct family of pygostylian avialans known from the Early Cretaceous, found in northern China. They are commonly placed as a sister group to Ornithothoraces, a group that contains all extant birds along with their clo ...
. The structure provided an evolutionary advantage, as a completely mobile tail as found in species such as ''
Archaeopteryx ''Archaeopteryx'' (; ), sometimes referred to by its German name, "" ( ''Primeval Bird'') is a genus of bird-like dinosaurs. The name derives from the ancient Greek (''archaîos''), meaning "ancient", and (''ptéryx''), meaning "feather" ...
'' is detrimental to its use for flight control. Modern birds still develop longer caudal vertebrae in their
embryo An embryo ( ) is the initial stage of development for a multicellular organism. In organisms that reproduce sexually, embryonic development is the part of the life cycle that begins just after fertilization of the female egg cell by the male sp ...
nic state, which later fuse to form a pygostyle. There are two main types of pygostyle: one, found in Confuciusornithidae,
Enantiornithes The Enantiornithes, also known as enantiornithines or enantiornitheans in literature, are a group of extinct Avialae, avialans ("birds" in the broad sense), the most abundant and diverse group known from the Mesozoic era. Almost all retained teet ...
, and some other
Mesozoic The Mesozoic Era is the Era (geology), era of Earth's Geologic time scale, geological history, lasting from about , comprising the Triassic, Jurassic and Cretaceous Period (geology), Periods. It is characterized by the dominance of archosaurian r ...
birds, as well as in some
oviraptorosaur Oviraptorosaurs ("egg thief lizards") are a group of feathered maniraptoran dinosaurs from the Cretaceous Period of what are now Asia and North America. They are distinct for their characteristically short, beaked, parrot-like skulls, with or wi ...
s like ''
Nomingia ''Nomingia'' is a genus of Oviraptorosauria, oviraptorosaur theropod dinosaur hailing from the Late Cretaceous Bugin Tsav Beds of Mongolia. Discovery and naming The remains, consisting of most of the vertebral column, pelvic girdle and left ...
'', is long and rod- or dagger blade-like. None of the known fossils with such pygostyles show traces of well-developed rectrices. The tail feathers in these animals consisted of downy fuzz and sometimes 2–4 central "streamers" such as those found in some specimens of ''Confuciusornis'' or in ''
Paraprotopteryx ''Paraprotopteryx'' is a genus of enantiornithean birds from the Mesozoic of China.Zheng, X. Zhang, Z. & Hou, L. (2007) A new enantiornithine bird with four long rectrices from the Early Cretaceous of northern Hebei, China. ''Acta Geologica Sini ...
''.Clarke ''et al.'' (2006) By contrast, the function of the pygostyle in the
terrestrial Terrestrial refers to things related to land or the planet Earth, as opposed to extraterrestrial. Terrestrial may also refer to: * Terrestrial animal, an animal that lives on land opposed to living in water, or sometimes an animal that lives on o ...
''Nomingia'' is not known. It is notable however that its older relative ''
Caudipteryx ''Caudipteryx'' (meaning "tail feather") is a genus of small oviraptorosaur dinosaurs that lived in Asia during the Early Cretaceous, around 124.6 million years ago. They were feathered and extremely birdlike in their overall appearance, to the ...
'' had no pygostyle but a "fan" of symmetrical feathers which were probably used in social display. Perhaps such ornaments were widespread in
Caenagnathoidea Oviraptorosaurs ("egg thief lizards") are a group of feathered maniraptoran dinosaurs from the Cretaceous Period of what are now Asia and North America. They are distinct for their characteristically short, beaked, parrot-like skulls, with or wi ...
and their relatives, and ultimately the oviraptorosaurian pygostyle evolved to help support them. The related ''
Similicaudipteryx ''Similicaudipteryx'', meaning "similar to ''Caudipteryx''" (which itself means "tail feather"), is a genus of theropod dinosaur of the family Caudipteridae. Discovery and naming In 2007 the find of a new caudipterid was reported. In 2008 the ...
'', described in 2008, also had a rod-like pygostyle, associated with a fan of tail feathers. The other pygostyle type is
plowshare In agriculture, a plowshare (Differences between American and British spellings, US) or ploughshare (Differences between American and British spellings, UK; ) is a component of a plow (or plough). It is the cutting or leading edge, preceding t ...
-shaped. It is found in
Ornithurae Ornithurae (meaning "bird tails" in Greek) is a natural group that includes modern birds and their very close relatives such as the ichthyornithines and the hesperornithines. This clade is defined in the ''PhyloCode'' by Juan Benito and collea ...
(living birds and their closest relatives), and in almost all flying species is associated with an array of well-developed rectrices used in maneuvering. The central pair of these attach directly to the pygostyle, just as in ''
Confuciusornis ''Confuciusornis'' is a genus of basal crow-sized avialan from the Early Cretaceous Period of the Yixian and Jiufotang Formations of China, dating from 125 to 120 million years ago. Like modern birds, ''Confuciusornis'' had a toothless beak, bu ...
''. The other rectrices of Ornithurae are held in place and moved by structures called ''bulbi rectricium'' (rectricial bulbs), a complex feature of fat and muscles located on either side of the pygostyle. The oldest known species with such a pygostyle is '' Hongshanornis longicresta''. As evidenced by the
oviraptorosauria Oviraptorosaurs ("egg thief lizards") are a group of feathered maniraptoran dinosaurs from the Cretaceous Period (geology), Period of what are now Asia and North America. They are distinct for their characteristically short, beaked, parrot-like s ...
n cases, the pygostyle evolved at least twice, and rod-shaped pygostyles seem to have evolved several times, in association with shortening of the tail but not necessarily with a retractable fan of tail feathers. In other words, the pygostyles of oviraptorosaurs and ''Confuciusornis'' were likely weight-saving measures, and the specialized "true" pygostyles of ornithurans were adapted from these later to improve flight performance. The bird
clade In biology, a clade (), also known as a Monophyly, monophyletic group or natural group, is a group of organisms that is composed of a common ancestor and all of its descendants. Clades are the fundamental unit of cladistics, a modern approach t ...
Pygostylia Pygostylia is a group of avialans which includes the Confuciusornithidae and all of the more advanced species, the Ornithothoraces. Definition The group Pygostylia was intended to encompass all avialans with a short, stubby tail, as opposed t ...
was named in 1996, by
Luis Chiappe Luis María Chiappe (born 18 June 1962) is an Argentine paleontologist born in Buenos Aires who is best known for his discovery of the first sauropod nesting sites in the badlands of Patagonia in 1997 and for his work on the origin and early evol ...
, for the presence of this feature and roughly corresponds to its appearance in the bird family tree, though the feature itself is not included in its definition. In 2001,
Jacques Gauthier Jacques Armand Gauthier (born June 7, 1948, in New York City) is an American vertebrate paleontologist, comparative morphologist, and systematist, and one of the founders of the use of cladistics in biology. Life and career Gauthier is the ...
and
Kevin de Queiroz Kevin de Queiroz is a vertebrate, Evolutionary biology, evolutionary, and Systematics, systematic biologist. He has worked in the phylogenetics and evolutionary biology of squamate reptiles, the development of a unified species concept and of a P ...
(2001) re-defined Pygostylia to refer specifically to the
apomorphy In phylogenetics, an apomorphy (or derived trait) is a novel Phenotypic trait, character or character state that has evolution, evolved from its ancestral form (or Plesiomorphy and symplesiomorphy, plesiomorphy). A synapomorphy is an apomorphy sh ...
of a short tail bearing an avian pygostyle.


Etymology

"Pygostyle" is formed from
Ancient Greek Ancient Greek (, ; ) includes the forms of the Greek language used in ancient Greece and the classical antiquity, ancient world from around 1500 BC to 300 BC. It is often roughly divided into the following periods: Mycenaean Greek (), Greek ...
words, literally meaning "rump pillar". The insulting equation of the pygostyle with the noses of dignitaries dates back to 18th century Britain where ''pope's nose'' first appears (in 1786). The form ''parson's nose'' appears much later, dated to 1839. The usage is somewhat dependent on either a Catholic or Protestant viewpoint. The forms ''bishop's nose'' and ''sultan's nose'' are 20th century variants.


As food

Turkey tail or turkey butt has an international exportation market in places such as
Micronesia Micronesia (, ) is a subregion of Oceania, consisting of approximately 2,000 small islands in the Northwestern Pacific Ocean. It has a close shared cultural history with three other island regions: Maritime Southeast Asia to the west, Poly ...
,
Samoa Samoa, officially the Independent State of Samoa and known until 1997 as Western Samoa, is an island country in Polynesia, part of Oceania, in the South Pacific Ocean. It consists of two main islands (Savai'i and Upolu), two smaller, inhabited ...
, and
Ghana Ghana, officially the Republic of Ghana, is a country in West Africa. It is situated along the Gulf of Guinea and the Atlantic Ocean to the south, and shares borders with Côte d’Ivoire to the west, Burkina Faso to the north, and Togo to t ...
. The turkey tail is commonly exported from America because it is considered unhealthy and cut off the normal turkey. After World War II, cheap imported turkey tails became popular in Samoa. Because the cut has a very high fat content, it was banned from 2007 to 2013 to combat obesity, only allowed back when Samoa joined the World Trade Organization. The meat is otherwise used in
pet food Pet food is animal feed intended for consumption by pets. Typically sold in pet stores and supermarkets, it is usually specific to the type of animal, such as dog food or cat food. Most meat used for animals is a byproduct of the human food indus ...
.


Footnotes


References

*
Electronic Appendix
* {{refend Bird anatomy Dinosaur anatomy Ancient Greek