Článek Revert the (runners, ostriches and relatives) is about more than promised article on this blog. Actually, I promise it so long that I do not remember (and I do not want to look for) when it happened the first time. Now comes the time in any event. The purpose of the article is to address the three fundamental questions: 1) internal příbuzenstvím between the runners and their relationship to all other birds, 2) their nelétavosti and 3) their role in the debate about the origin of birds themselves. In addition, a brief plan to include other issues such as eg biogeography. And because the materials from which to draw, there is a wealth of, probably would be better if instead of a single article will set a new series.
first Nomenclature
It is probably advisable to get rid of this issue at the beginning, when reading further complicate life. Ratitae is a group that traditionally includes a number of flightless birds living in the southern hemisphere continents, usually possessing huge physical dimensions. These include Ostrich, rheas, emus, cassowaries, kiwi and a few extinct taxa (birds moa, "elephant bird" Aepyornis ). Name Ratitae kurzivuji because it bears a phylogenetic definition (Gauthier & de Queiroz 2001), which, however, is a prime example of how to define nothing. In some phylogenetic hypothesis does not contain anything other than the two-ostrich; else again almost all non-passeriformní birds. Because part of this article will be debate among others about whether the ostrich, cassowaries, kiwi, emu and rhea are really natural (monophyletic) group, to which no other living bird is not, is phylogenetically defined name even useless - its definition is simple text monophyly requires change can only content. Here, by contrast, need for fixed content; about whether the group is made up of representatives of traditional and Revert nothing monophyletic, paraphyletic or polyphyletic, remain to be discuss. And as you probably no ornithologist when they hear the name Ratitae equips Gauthier and de Queirozovu definition, allow her to ignore the text.
Czech Revert is the name of "runners". But it is now also used this name for a group Palaeognathae , which in addition to Revert as such includes even tinamy. (A relative of Tina and Revert still be talking about.) You'll concepts of "runners" and " Ratitae / Revert" to use as a free interchangeable synonyms.
in ornithological literature, more than Ratitae Struthioniformes uses an alternative name, perhaps because it is podnější names of the other Birds orders (Galliformes, Passeriformes ,...). Its scope is not clear: sometimes the same range as Ratitae (eg Sibley & Ahlquist 1981), is sometimes limited to the two-ostrich (Struthio camelus ) and the other runners are promoted to their own 'laws'. Rhea (Rhea ) became the sole representative of the "order" Rheiformes, cassowaries and get emuové "order" Casuariformes etc. This approach, however preposterously inflated taxonomy: whether you say 'rhea (Rhea ), "nanduovití ( Rheidae) or "nanduové (Rheiformes), we always mean the same thing: about 8 distinct taxa seskupovaných into" types " Rhea americana (rhea Rhea) and Rhea pennata (Darwin's rhea). The creation of such names without substance apparently stems from a desire to make everything, what was in ornithological taxonomy're sure to fall in the "relationship between the Council" (interordinal Relationships). For these reasons, I use Struthioniformes: Unlike Ratitae book one can not be sure what to expect from him.
Living runners ( Ratitae , Struthioniformes sensu lato). From the position of top left clockwise: brown emu (Dromaius novaehollandiae ), Ostrich ( Struthio camelus), rhea Rhea (Rhea americana ), Southern Cassowary (Casuarius casuarius ), kiwi? South (Apteryx ? australis). (Sources: author's own photographs of Prague zoo [emu, ostrich - Spring 2010, rhea - Winter 2011], cassowaries - modified on jennifermarohasy.com, kiwi - modified from the info-rotorua.com)
second Inspiration
Runners are undoubtedly one of the strangest and most interesting groups of birds. The debate about the origin of birds, which are fully erupted immediately after the publication of Darwin's book The Origin of Species (Darwin 1859), played a prominent role. Flightless birds are plenty, but few of them that has caught the bird's body's modification of the plan so far as they are. This is obviously all very interesting, but what made me write something for runners, was a high school textbook Fundamentals of Biology (Kislinger et al. 1995). These are
many respects, a truly remarkable publications. Although the piece that I got their hands on enough pages had yellowed out to get the look of something that lay buried for fifty years somewhere on the land are Basics far, far younger - without at least one of reflected. When leafing through a man encounters with such pearls, which quickly led me to the shame that I am imperfect phylogeny of publications criticizing type amazing world of dinosaurs . Well, on purpose, knowing you know that "mushrooms" (meant the sea, which is usually called Sponges) evolved from the "element" already 2.5 billion years ago (!) and that "this line was blank at the end, even though its representatives living today? One would then almost did not recognize her from the line, which ends blindly ... You will find everything, including romerogramu where birds, Pterosaur, crocodiles and dinosaurs independently splits off from the "thekodontů (Fig. 32). Dvojdyšní, scaly and hatérie have the unenviable fate of the same romerogramu, designated as an "evolution of vertebrates, is not fit. There are also scripts Brontosaurus (Panebože! really this could mean that anyone?), Tail pulling the country and representing the group "veleještěrů. As far as birds and runners, we read that:
many respects, a truly remarkable publications. Although the piece that I got their hands on enough pages had yellowed out to get the look of something that lay buried for fifty years somewhere on the land are Basics far, far younger - without at least one of reflected. When leafing through a man encounters with such pearls, which quickly led me to the shame that I am imperfect phylogeny of publications criticizing type amazing world of dinosaurs . Well, on purpose, knowing you know that "mushrooms" (meant the sea, which is usually called Sponges) evolved from the "element" already 2.5 billion years ago (!) and that "this line was blank at the end, even though its representatives living today? One would then almost did not recognize her from the line, which ends blindly ... You will find everything, including romerogramu where birds, Pterosaur, crocodiles and dinosaurs independently splits off from the "thekodontů (Fig. 32). Dvojdyšní, scaly and hatérie have the unenviable fate of the same romerogramu, designated as an "evolution of vertebrates, is not fit. There are also scripts Brontosaurus (Panebože! really this could mean that anyone?), Tail pulling the country and representing the group "veleještěrů. As far as birds and runners, we read that:
-----> relatives original small dinosaurs are birds (the jury)
- [...]
runners are individually spin-off from small dinosaurs. Do not fly!
Kislinger et al. 1995:120
Wow. Let's see. The authors had to be very sure about when it submit to a cramming school students - there is an exclamation point, a. .. and all. No wonder that in such a reference will cause an enormous interest in the history of research of the runners, if you know of a single case where a similar hypothesis was proposed, with the author if any reasonable ornithologist laughed. What you mean, it will be apparent from the subsequent part of the article.
third History
Have Linnaeus (1758) knew the three representatives of the runners when the 10th edition of his Systema Naturae publications founded the modern biological nomenclature. Named after the Struthio camelus, Struthio casuarius and Struthio americanus. This first (the two-ostrich) are known by that name yet. The second taxon was two years later renamed Brisson Casuarius casuarius (casuar) and the third time to become Rhea americana or rhea. Another discovery quickly followed, when Latham in 1790 described a "casuar novoholandského. Around the same time, paradoxically, in New Zealand last remnants of dying birds moa (Dinornithidae), as up to the largest birds, the world has seen since the Upper Cretaceous oviraptorosaurů. Shaw (1813) from New Zealand described kiviho, the smallest and perhaps even nejpodivnějšího Revert, and three years later was "cassowaries novoholandský" renamed to emus (Dromaius taxon ). Then the foundations scheme runners received more or less normal appearance.
3.1 Identifying runners
first person to recognize that the ostrich, emu, rheas, cassowaries are among the birds living together was probably Merrem (1813). The classification of birds divided into two groups: Ratitae include the above-mentioned taxa (kiwi Merremovi not yet known, early on it was, however, included), Carinatae all other birds, characterized by a keel on the sternum for the insertion of swing muscles (which need flightless Revert). Lesson (1831) Merremovy Revert kiviho took over and instituted a new "family" name Nullipennes .* Both groups Ratitae Nullipennes and then summarized the "anormaux oiseaux" ("abnormal birds"). The revolution here - as in many other areas - came with Darwin (1859), which among others has shown that all the previous classification, often unconsciously, trying to approximate a genealogy, a common origin. Already in the first edition The Origin of Species Darwin runners at several points mentioned: kiviho called vestigial wings (Darwin 2007:508) and thus indicated that his ancestors were Glider same way as most other birds. This hypothesis is supported by, paradoxically, one of the largest and most influential opponents of Darwin, the comparative anatomist Richard Owen (who also introduced the name Dinosauria ). Owen, while protesting that the main mechanism of evolution should be a natural choice, with a common origin, but not a problem. It is therefore not surprising that actively involved in discussions about ancestors birds (Owen 1875) and also the debate about the origin and kinship Revert.
3.2 Owen and moa birds
At this point it is worth to note that Owen, in previous years, his research and taxonomy of birds moa. The fact that New Zealand had been inhabited by giant Revert, Owen persuaded a fragment of femur and the author of this finding then published (Owen 1839). Few of us are convinced, however, many scientists believed that the Pacific concentrate was too small to feed the population of flightless birds with similar dimensions. Owen have started to collect more bones and systematically compared with ostriches, emu, rhea and kiwi. He found that, like the kiwi - but unlike all other Revert - no moa at the femoral neck air vent, and that the body was so filled with bone marrow rather than pneumatizované. Kivimu MOA is similar in the fact that tarzometatarzus (feet) was only half as long as tibiotarzus; tibiotarzu contrast ratio of length to the femur (2-1) answered rather different ratitům (Anderson 1989). Four years after it was Owen with moa bird skeletal material for the first time confronted him definitively described under the name "Megalornis" Nova Zealandiae .** Very soon (description was still in print) has proven that prenomen Megalornis already absorbed, and Owen then changed the formal name for the moa Dinornis .
Systematics moa birds, however, quickly began to unravel. From the description D. novaezealandiae passed only a year when the first bird moa followed by a second, D. giganteus (Moa Island in the South, Owen 1844), and a few months (Owen 1844b) later and the third ( D. struthoides ), fourth (D didiformis ), fifth ( D. dromaeoides ) and sixth ( D. otidiformis ). For a complete list of all subtaxa dinornise present Worthy & Holdaway (2002), of the table shows that in the late 50th of the 19 century, the taxonomy dinornitidů confusing mess. Far more interesting, however, was the development of views on MOA relationship to other birds. Anderson (1989) states that the findings of more complete skeletons Owen was increasingly tempted to withdraw their initial conclusions about the nature dinornitidů runners. In particular, a combination of lack of a fourth toe, and beak-shaped pans and "crocodile cranium" led Owen (1848) to speculate about their close relationship with the drop (Otididae), which refused to end just because the shape of the front of the beak. In the same work even Owen (1848) described the taxon Palapteryx (now generally regarded as synonymous dinornise) for Revert (strutionida ") transition between kiwis and emus, while Dinornis have anatomy" so strange that the author does not assign gender to any known natural family of birds. Its location in order Struthionidae suggest anything more than deadlock development of wings and exaggerated foot development, arranged for movement on dry land. " (Owen 1848:8) About the same time conducted a heated debate about how the moa birds appeared. Owen (1844b) pointed out that the combined proportion of short and robust tarzometatarzu cervical vertebrae resembled rather than casuar kiviho or African ostrich, and even casuar chosen as a model, by which time the moa bird reconstructed:
Worthy & Holdaway (2002) notes, however, that Owen seems to live casuar never seen before - otherwise it would have to know that the body is oriented horizontally much more than it is in its reconstruction. The same authors note that, paradoxically, Owen was right - cassowaries indeed of all living birds, the moa Revert like most, it's just that the normal, relaxed position looks very different than the illustration Owen. The front part synsakra (typically bird bones, resulting from adhesions between the pelvic bones, and sacral vertebrae) is parallel to the surface level, and thoracic vertebrae as well - this is even for all living except Revert kiviho (Worthy & Holdaway 2002:163).
Discussions also led to whether the moa birds have wings at all. It is a well known fact that all living Runners have wings, even compared létavým dwarf birds. With a little effort would go even build a sequence leading from the ostrich, whose wings are still prominent and actively used (for stabilization at or during the course of the run), up to kivimu for which there are almost no visible wings and 13 squadrons are reduced to the extent that - as aptly noted Matt Martyniuk - kompsognátidů resemble feathers. Where in this sequence fits me? Owen had already in 1843 noticed that the whole collection missing front leg bones - but because he knew that the absence of evidence is not evidence of absence, decided to use "argument by homology. Homology is a concept which is still recovering from biology předevolučních Ever meant something entirely different than today (which is essentially used interchangeably with the term "synapomorfie). Owen assumed that there is a relationship between the degree of pneumatization of the skeleton and the size of the wings. The ostrich and rhea, who are living on the wing developed best runners are pneumatizovány skull bones, spine, ribs, pelvis and sternum, femur, and ravens (Anderson 1989). Emus and cassowaries have smaller wings, air sacs still interfere with the femur. For kiviho air bags do not infringe on the body than the chest. If the relationship waylaid applied, it should have wings MOA between the state transition observed in kiviho and emus: Dinornis showed otvůrky Although the air sacs (pneumatic foramen) on the vertebrae, but not in the femur. The hypothesis seemed confirmed when in 1850, Owen received a fragment of an alleged humerus (arm bone). The ultimate solution to the issue appeared to dinornise wings in 1864. At Mo's was discovered skapulakorakoid (shoulder girdle), which did not carry the slightest sign of the glenoid: hole through which the scapula and humerus joints thus the entire forelimb. Anderson (1989) cites Owen, that in itself would not result complete absence of wings - but it would require a bizarre situation where the wing joined to the rest of the body only ligaments Owen (1866 :170-171) alone and rejected instead, it opined skapulakorakoid that the skeleton remained only as a lever for the respiratory muscles. Although
moa birds should go take a long time (only the beginning of the 20th century, the taxonomic history has grown so much that all details you can actually afford to go just a book) and have them in series will return - as the occasion Cracraftových systematic revisions - the sub-heading I will conclude with the first hypothesis of paleogeography. In the mid 19th century, of course, not known to move continents, and therefore all the more surprising that it was the Revert and among them particularly birds Moa, who already at that time caused some scientists to speculate about such mechanisms. Owen initially speculated about the "mighty wave unstable and constantly shifting the earth's crust, which has entered the remains dinornitidům similar birds throughout the Pacific to North America and the sole remnant should be just New Zealand (Anderson, 1989:41). He later left the speculation, with the ancestors of birds Moa in New Zealand just flew (Owen 1879 - on the same site by the way the author criticizes and rejects the hypothesis of the origin of birds Saurian) before, however, inspired several authors to the hypothesis about the entire southern continent, very similar to what we now call Gondwana. For the first time it probably gave Hooker (1853) and elaborated in detail by Forbes (1893). According to him, was once the southern hemisphere oceans, all ground to a depth of 3.7 km land, which connected Africa, Antarctica and South America into a single supercontinent. Although this view proved to be correct (and the geographical distribution of key runners), in its time course, such support did not record. Forbes also believed that ratites including dinornitidů represent a natural group, in its classification of them, however, has not such an exceptional place as Merry and grouped them together with hrabavými (McDowell 1948).
3.3 Views on the origin and kinship in the years 1859-1868 Revert
It was probably just Owen's research on bird moa, which it in 1866 inspired an unusual hypothesis. In their classification, all the while runners into one group, as was the custom Merremových times, but also by the view according to which the ostrich (Struthio ) more closely related dropovitým (Otididae) than the rest of the group, while the MOA ( Dinornis ) and kiwi (Apteryx ) are closer to tabonovitým (Megapodidae). Thus formulated the hypothesis that a total of bizarre, but otherwise anticipates "Revert to fight", which will continue until 21 century. If are indeed the main characteristic of runners stunted wings and nelétavost, as he wrote Owen (1866b), what prevents these characters gain various groups of birds in a long time span independently of each other?
Huxley, who - unlike Owen - inclined to Darwin's theory of evolution immediately after its publication, had the whole thing exactly the opposite view. Living Revert regarded as a remnant of ancient, once-great groups of birds, who lost years, but never becomes. Until then it occurred Glider, karinátní birds. In January 1868 letter to Huxley in the German evolutionary biologist Ernst Haeckel wrote:
Systematics moa birds, however, quickly began to unravel. From the description D. novaezealandiae passed only a year when the first bird moa followed by a second, D. giganteus (Moa Island in the South, Owen 1844), and a few months (Owen 1844b) later and the third ( D. struthoides ), fourth (D didiformis ), fifth ( D. dromaeoides ) and sixth ( D. otidiformis ). For a complete list of all subtaxa dinornise present Worthy & Holdaway (2002), of the table shows that in the late 50th of the 19 century, the taxonomy dinornitidů confusing mess. Far more interesting, however, was the development of views on MOA relationship to other birds. Anderson (1989) states that the findings of more complete skeletons Owen was increasingly tempted to withdraw their initial conclusions about the nature dinornitidů runners. In particular, a combination of lack of a fourth toe, and beak-shaped pans and "crocodile cranium" led Owen (1848) to speculate about their close relationship with the drop (Otididae), which refused to end just because the shape of the front of the beak. In the same work even Owen (1848) described the taxon Palapteryx (now generally regarded as synonymous dinornise) for Revert (strutionida ") transition between kiwis and emus, while Dinornis have anatomy" so strange that the author does not assign gender to any known natural family of birds. Its location in order Struthionidae suggest anything more than deadlock development of wings and exaggerated foot development, arranged for movement on dry land. " (Owen 1848:8) About the same time conducted a heated debate about how the moa birds appeared. Owen (1844b) pointed out that the combined proportion of short and robust tarzometatarzu cervical vertebrae resembled rather than casuar kiviho or African ostrich, and even casuar chosen as a model, by which time the moa bird reconstructed:
first reconstruction Owen Bird MOA. The assumption that his posture resembled living cassowaries (pictured left), Owen concluded the height about 10 feet (about 3 meters). The illustrations, however, that much of the possession kasuářím body did not know ... (Source: Owen 1844: Plate XXX digitized here )
Worthy & Holdaway (2002) notes, however, that Owen seems to live casuar never seen before - otherwise it would have to know that the body is oriented horizontally much more than it is in its reconstruction. The same authors note that, paradoxically, Owen was right - cassowaries indeed of all living birds, the moa Revert like most, it's just that the normal, relaxed position looks very different than the illustration Owen. The front part synsakra (typically bird bones, resulting from adhesions between the pelvic bones, and sacral vertebrae) is parallel to the surface level, and thoracic vertebrae as well - this is even for all living except Revert kiviho (Worthy & Holdaway 2002:163).
Discussions also led to whether the moa birds have wings at all. It is a well known fact that all living Runners have wings, even compared létavým dwarf birds. With a little effort would go even build a sequence leading from the ostrich, whose wings are still prominent and actively used (for stabilization at or during the course of the run), up to kivimu for which there are almost no visible wings and 13 squadrons are reduced to the extent that - as aptly noted Matt Martyniuk - kompsognátidů resemble feathers. Where in this sequence fits me? Owen had already in 1843 noticed that the whole collection missing front leg bones - but because he knew that the absence of evidence is not evidence of absence, decided to use "argument by homology. Homology is a concept which is still recovering from biology předevolučních Ever meant something entirely different than today (which is essentially used interchangeably with the term "synapomorfie). Owen assumed that there is a relationship between the degree of pneumatization of the skeleton and the size of the wings. The ostrich and rhea, who are living on the wing developed best runners are pneumatizovány skull bones, spine, ribs, pelvis and sternum, femur, and ravens (Anderson 1989). Emus and cassowaries have smaller wings, air sacs still interfere with the femur. For kiviho air bags do not infringe on the body than the chest. If the relationship waylaid applied, it should have wings MOA between the state transition observed in kiviho and emus: Dinornis showed otvůrky Although the air sacs (pneumatic foramen) on the vertebrae, but not in the femur. The hypothesis seemed confirmed when in 1850, Owen received a fragment of an alleged humerus (arm bone). The ultimate solution to the issue appeared to dinornise wings in 1864. At Mo's was discovered skapulakorakoid (shoulder girdle), which did not carry the slightest sign of the glenoid: hole through which the scapula and humerus joints thus the entire forelimb. Anderson (1989) cites Owen, that in itself would not result complete absence of wings - but it would require a bizarre situation where the wing joined to the rest of the body only ligaments Owen (1866 :170-171) alone and rejected instead, it opined skapulakorakoid that the skeleton remained only as a lever for the respiratory muscles. Although
moa birds should go take a long time (only the beginning of the 20th century, the taxonomic history has grown so much that all details you can actually afford to go just a book) and have them in series will return - as the occasion Cracraftových systematic revisions - the sub-heading I will conclude with the first hypothesis of paleogeography. In the mid 19th century, of course, not known to move continents, and therefore all the more surprising that it was the Revert and among them particularly birds Moa, who already at that time caused some scientists to speculate about such mechanisms. Owen initially speculated about the "mighty wave unstable and constantly shifting the earth's crust, which has entered the remains dinornitidům similar birds throughout the Pacific to North America and the sole remnant should be just New Zealand (Anderson, 1989:41). He later left the speculation, with the ancestors of birds Moa in New Zealand just flew (Owen 1879 - on the same site by the way the author criticizes and rejects the hypothesis of the origin of birds Saurian) before, however, inspired several authors to the hypothesis about the entire southern continent, very similar to what we now call Gondwana. For the first time it probably gave Hooker (1853) and elaborated in detail by Forbes (1893). According to him, was once the southern hemisphere oceans, all ground to a depth of 3.7 km land, which connected Africa, Antarctica and South America into a single supercontinent. Although this view proved to be correct (and the geographical distribution of key runners), in its time course, such support did not record. Forbes also believed that ratites including dinornitidů represent a natural group, in its classification of them, however, has not such an exceptional place as Merry and grouped them together with hrabavými (McDowell 1948).
3.3 Views on the origin and kinship in the years 1859-1868 Revert
It was probably just Owen's research on bird moa, which it in 1866 inspired an unusual hypothesis. In their classification, all the while runners into one group, as was the custom Merremových times, but also by the view according to which the ostrich (Struthio ) more closely related dropovitým (Otididae) than the rest of the group, while the MOA ( Dinornis ) and kiwi (Apteryx ) are closer to tabonovitým (Megapodidae). Thus formulated the hypothesis that a total of bizarre, but otherwise anticipates "Revert to fight", which will continue until 21 century. If are indeed the main characteristic of runners stunted wings and nelétavost, as he wrote Owen (1866b), what prevents these characters gain various groups of birds in a long time span independently of each other?
Huxley, who - unlike Owen - inclined to Darwin's theory of evolution immediately after its publication, had the whole thing exactly the opposite view. Living Revert regarded as a remnant of ancient, once-great groups of birds, who lost years, but never becomes. Until then it occurred Glider, karinátní birds. In January 1868 letter to Huxley in the German evolutionary biologist Ernst Haeckel wrote:
In scientific work the main thing just now about Which I Am Engaged with a revision of the Dinosauria - with an eye to the Descendenz Theorie ! The Road from Reptiles to Birds is the way of Dinosauria to the Ratitae - The Bird 'phylum' WAS Struthious, and Wings Grew out of Fore rudimentary limbs. You See That Among Other Things I have been reading Ernst Haeckel's Morphologie .
TH Huxley, quoted in Switek 2010:255
Huxley therefore considered that the front legs of runners representing the "atrophied" remains of a once functional wings, as proposed by Owen - ostriches and their relatives have in this form to inherit from the dinosaur (or dinosaur like) ancestors. At that time, but Archaeopteryx and anatomical analysis of the various authors show that the whole scenario looked different. Bavarian "prapták" was in many respects more primitive than runners (Teeth in the jaws, a long tail bone), and yet had no wings létavými comparable to modern birds. Huxley - Darwin as well - but he showed a great interest and archeopteryga his hypothesis ratitních ancestors of birds and spoil the future. As shown Switek (2010), in the author's idea of \u200b\u200bthe runners have evolved from small dinosaurs svrchnojurského kompsognáta type (which Huxley later removed from dinosaurs, erected for him taxon "Compsognatha at the same level in both dinosaurs and the summarized group taxon Ornithoscelida) and its extension "Rudimentary forelegs" gave rise to karinátům. Huxley (1868) expressed their distrust archeopteryga the following sentence:
In many RESPECTS, Archaeopteryx is more remote from the boundary-line Between Birds and Reptiles Than Some living Ratitae are.
Huxley 1868:248
Darwin's theory was launched at the same time other research programs, including the creation of new classifications, which reflect as closely as possible genealogical relationships. Parker (1864) as Revert to support monophyly, based on pterylografie (Organization of feathers). In the same work as the first author advocated a close relationship between the runners and South American tinamami. Previously, these birds - who, unlike runners and fly to the keel of the sternum are - associated more with hrabavými (Lesson 1831). Parker deeply ahead of its time: Taxon connecting the runner and is now called tinamy Palaeognathae . Rather
fad phylogenetic hypothesis represented by a short Note outlined Cope (1867). According to the origin of runners (and / or penguins) has nothing to do with wild birds and is instead primarily flightless descendants of dinosaurs.
Parker Revert spent on research (see eg Parker 1866), his conclusions about their kinship tinamám however, overshadowed the first explicit evolutionary classification of birds, stood again for the Huxley (1867). He, like Parker noted the similarities between the bone floor runners and Tina - Anatomical features of the floor was all based his classification of birds, and the type shared between these two groups Huxley called dromeognátní (dromaeognathous palate). As described Gussekloo (2000), the original definition of the character and include) the absence of contact between the rear end of the bone plate and parasfenoidním rostro (= rostroparasfenoidem, kultriformním process - see above illustration), b) the absence of contact between the front end of the wing bone (pterygoid) and parasfenoidním rostro c) strong bazipterygoidní salient. Over time, the definition and modified, and some authors even the very existence of a specific paleognátního or "dromeognátního" floor rejected due to excessive diversity that this element of the runners reported (McDowell 1948 for a discussion about the validity of letters, see also Gussekloo & Zweers 1999). Correspondence between dromeognátním floor runners and Tina stood Huxley, in any event prior to the problem: while this character combining both groups, the keel of the sternum and the ability to fly to the contrary, it included tinamy Merremových karinátů. The author finally decided to respect the traditional division and its classification introduced three "sub": Saururae archeopterygem as the sole representative Ratitae with traditional ingredients and Carinatae covering all remaining birds. There is evident, inter alia, that the Huxley kept the back door of his hypothesis on the origin of birds ratitním Most other authors would probably not hesitate to connect with Karin Revert to one group to another archeopteryga sit, as it did before Huxley Haeckel (1866).
fad phylogenetic hypothesis represented by a short Note outlined Cope (1867). According to the origin of runners (and / or penguins) has nothing to do with wild birds and is instead primarily flightless descendants of dinosaurs.
Typically paleognátní bony palate (palatal-pterygoid complex "sensu Gussekloo) illustrated the interpretive drawing of skull rhea when viewed from below. (A) mouldboard bone, (B) palatine bone, (C) rostrum parasfenoidní / kultriformní the neck (D) bazipterygoidní ness (E) wing bone (F) bone square. (Source: Gussekloo & Bout, 2005: Figure 2)
the same photograph of the skull excised the two-ostrich (Struthio camelus ) . It is a famous female ostrich Veronica, who - after a happy reunion with Mike'm Taylor - featured in a series of posts on SV-POW, whence comes this photo and where you can see and its 3D version. Note the broad bone radličných that prevent reconciliation and pterygoid palatal bone with parasfenoidním rostro. (Source: svpow.wordpress.com)
Parker Revert spent on research (see eg Parker 1866), his conclusions about their kinship tinamám however, overshadowed the first explicit evolutionary classification of birds, stood again for the Huxley (1867). He, like Parker noted the similarities between the bone floor runners and Tina - Anatomical features of the floor was all based his classification of birds, and the type shared between these two groups Huxley called dromeognátní (dromaeognathous palate). As described Gussekloo (2000), the original definition of the character and include) the absence of contact between the rear end of the bone plate and parasfenoidním rostro (= rostroparasfenoidem, kultriformním process - see above illustration), b) the absence of contact between the front end of the wing bone (pterygoid) and parasfenoidním rostro c) strong bazipterygoidní salient. Over time, the definition and modified, and some authors even the very existence of a specific paleognátního or "dromeognátního" floor rejected due to excessive diversity that this element of the runners reported (McDowell 1948 for a discussion about the validity of letters, see also Gussekloo & Zweers 1999). Correspondence between dromeognátním floor runners and Tina stood Huxley, in any event prior to the problem: while this character combining both groups, the keel of the sternum and the ability to fly to the contrary, it included tinamy Merremových karinátů. The author finally decided to respect the traditional division and its classification introduced three "sub": Saururae archeopterygem as the sole representative Ratitae with traditional ingredients and Carinatae covering all remaining birds. There is evident, inter alia, that the Huxley kept the back door of his hypothesis on the origin of birds ratitním Most other authors would probably not hesitate to connect with Karin Revert to one group to another archeopteryga sit, as it did before Huxley Haeckel (1866).
is most appreciated in retrospect Huxley's classification karinátů, consisting in detecting four different types of bony palate, which is reflected in it as "advice" Dromaeognathae, Schizognathae, and Desmognathae Aegithognathae. Tinamy figure at the beginning karinátů as the only representatives of the "Order" and he Dromaeognathae Huxley (1867:426) recognized that "the bridge struthious Carinae of all birds". In the Revert Huxley recognized the five unnamed groups: ostrich, rhea, kiwi and moa and had four of them for themselves (which shows that Huxley's arguments failed to convince Owen of kinship kiviho and dinornitidů). Emus and cassowaries, then shared a fifth group. As the show's future, this volume remains the only lasting security in a changing taxonomy Revert. Huxley's classification of birds in any event, opens a new era: the Sibley & Ahlquist (1972) showed that in the history of avian systematics since it runs clear line drawn across Fürbringera, Gadowa and Wetmore to today's "default" in the various classifications Checklist.
* standardized suffix "-idaea 'for all zoological names" at the family " introduced to the so-called stricklandský code in 1843, for an overview of the historical development level nomenclature see de Queiroz (2005). ** Sample
similar phenomenon - although today it is a fundamental rule of zoological nomenclature, the "generic" name must consist of a single word without hyphens and accents, in the 1 19th century such universal agreement even existed.
Sources:
* standardized suffix "-idaea 'for all zoological names" at the family " introduced to the so-called stricklandský code in 1843, for an overview of the historical development level nomenclature see de Queiroz (2005). ** Sample
similar phenomenon - although today it is a fundamental rule of zoological nomenclature, the "generic" name must consist of a single word without hyphens and accents, in the 1 19th century such universal agreement even existed.
Sources:
- Anderson, AJ 1989 [2003] Prodigious Birds: Moas and Moa-Hunting in Prehistoric New Zealand . Cambridge: Cambridge Univ Press. 260 p
- Cope ED 1867 An account of extinct reptiles which approached the birds. Proc Acad Nat Sci Philad 19: 234–5
- Darwin CR 1859 On the origin of species by means of natural selection, or the preservation of favoured races in the struggle for life . London: John Murray
- Darwin CR 1872[2007] [ O vzniku druhů přírodním výběrem .] Prague: Academia [Transl. by Hadač E, Hadačová A]
- de Queiroz K 2005 Linnaean, rank-based, and phylogenetic nomenclature: Restoring primacy to the link between names and taxa. Symb Bot Ups 33(3): 127–40
- Forbes HO 1893 The Chatham Islands: their relation to a former continent. Royal Geogr Soc Suppl Pap 3, Part 4
- Gauthier JA, de Queiroz K 2001 Feathered dinosaurs, flying dinosaurs, crown dinosaurs, and the name "Aves". 7–41 in Gauthier JA, Gall LF, eds, New Perspectives on the Origin and Early Evolution of Birds: Proceedings of the International Symposium in Honor of John H. Ostrom . New Haven: Peabody Mus Nat Hist, Yale Univ
- Gussekloo SWS 2000 The evolution of the palaeognathous birds: Functional morphology and evolutionary patterns. PhD thesis, Leiden Univ, Leiden. 182 p
- Gussekloo SWS, Bout RG 2005 Cranial kinesis in palaeognathous birds. J Exp Biol 208: 3409–19
- Gussekloo SWS, Zweers GA 1999 The paleognathous pterygoid–palatinum complex. A true character? Neth J Zool 49: 29–43
- Haeckel E 1866 Generelle Morphologie der Organismen . Berlin: Georg Reimer. 462 p
- Hooker JD 1853 The Botany of the Antarctic Voyage of HM Discovery Ships Erebus and Terror in the Years 1839–1843, vol. II: Flora Novae Zelandiae . London: Lovell Reeve
- Huxley TH 1867 On the classification of birds; and on the taxonomic value of the modifications of certain of the cranial bones observable in that class. Proc Zool Soc Lond 1867: 415–72
- Huxley TH 1868 Remarks upon Archaeopteryx lithographica . Proc R Soc London 16: 243–8
- Kislinger F, Laníková J, Šlégl J, Žurková I 1995 Biologie V: Základy obecné biologie . Klatovy: Gymnázium Klatovy
- Lesson RP 1831 Traite d'ornithologie. Vol. 1 . Paris: Levrault. 659 p
- Linné C 1758 Systema naturae per regna tria naturae, secundum classes, ordines, genera, species, cum characteribus, differentiis, synonimis, locis. Tomus I. Editio decima, reformata . Stockholm: Laurentius Salvius. 824 p
- McDowell S 1948 The bony palate of birds. Part I. The Palaeognathae. Auk 65(4): 520–49
- Merrem B 1813 Tentamen systematis naturalis avium. Konigl. [Preussische] Akad Wiss Berlin, Abh 1812–13 [1816]: 237–59
- Owen R 1839 On the bone of an unknown Struthious bird from New Zealand. Proc Zool Soc Lond 1839: 169–70
- Owen R 1844a [Part 2 of Owen's memoir on Dinornis , read November 28, 1843.] Proc Zool Soc Lond 11(129): 144–6
- Owen R 1844b On Dinornis , an extinct genus of tridactyle struthious birds, with descriptions of portions of the skeleton of five species which formerly existed in New Zealand. Trans Zool Soc Lond 3(3): 243–76
- Owen R 1848 [April 13 – read 11 January 1848] On the remains of the gigantic and presumed extinct wingless or terrestrial birds of New Zealand ( Dinornis and Palapteryx ), with indications of two other genera( Notornis and Nestor ). Proc Zool Soc Lond 16(180): 1–11
- Owen R 1866a On Dinornis (Part IX): containing a description of the skull, atlas, and scapulocoracoid bone of the Dinornis robustus Owen. Trans Zool Soc Lond 5: 337–58
- Owen R 1866b Birds and mammals. On the anatomy of vertebrates, vol. 2 . London: Longmans, Green. 592 p
- Owen R 1875 Monograph of the fossil reptiles of the Liassic formations II. Pterosauria. Palaeontol Soc Monogr 41–81
- Owen R 1879 Memoirs on the extinct wingless birds of New Zealand; with an appendix on those in England, Australia, Newfoundland, Mauritius and Rodriguez . London: Van Voorst. 2 vols, 612 p
- Parker WK 1864 On the osteology of gallinaceous birds and tinamous. Trans Zool Soc Lond 5(3): 149–241
- Parker WK 1866 On the structure and development of the skull in the Ostrich Tribe. Phil Trans R Soc Lond 156: 113–83
- Shaw GA 1813 The Naturalist's Miscellany 24, pl. 1057, 1058
- Sibley CG, Ahlquist JE1972 A comparative study of egg-white proteins of non-passerine birds. Bull Peabody Mus Nat Hist 39: 1–276
- Sibley CG, Ahlquist JE 1981 The phylogeny and relationships of the ratite birds as indicated by DNA-DNA hybridization. 301–35 in Scudder GGE, Reveal JL, eds, Evolution Today, Proc 2nd Int Congr Syst Evol Biol . Pennsylvania: Pittsburgh
- Switek B 2010 Thomas Henry Huxley and the reptile to bird transition. Geol Soc Lond Spec Pub 343(1): 251–63
- Worthy TH, Holdaway RN 2002 The Lost World of the Moa: Prehistoric Life of New Zealand . Bloomington: Indiana Univ Press
0 comments:
Post a Comment