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Hesperornis a carnivorous marine ostrich difylie birds and the beginnings of the Wars of the runners monophyly (Ratitae, part 2)

In previous work was briefly zrekapitulována Revert history research since 1758, the beginning of modern zoological nomenclature *, Huxley breakthrough in the classification of birds from 1867. Since the theory of evolution was formulated just before the end of this interval was not so much about what to write: what was the point to ask whether any group of "natural" and whether they are runners Glider primarily or secondary, when he was the transformation of organisms at the time generally known fact ?

3.4 Systematics Revert between Huxley and Fürbringerem

Huxley's classification, dividing birds into three "subclasses" ( Carinatae , Ratitae , Saururae) and shlukující tinamy along with other flying Revert birds instead, was only the beginning of the dispute between the runners instead of the other birds. One of the first Huxley successor was Garrod (1874), in which we would classify along with that Huxley found much common ground. Garrod ignored extinct birds, including birds and archeopteryga MOA, then living among the group recognized the dichotomy Homalogonatae and Anomalogonatae. Anomalogonáti included only a singer and woodpeckers (United obojakého absence of M. ambiens muscle, allowing the finger grip after landing on a branch), then the remainder homalogonáti birds, divided into four orders. Garrod saw the runners as monophyletic, in contrast to previous authors, believing the uniqueness of this group, but they did not even own "order". It is summarized with hrabavými (Gallinaceae ") to" order Galliformes. Unexpectedly advanced feature Garrodovy scheme was that it merged with runners tinamami. While this has suggested Parker (1864, see last part of the article), but Garrod was the first to formally recognize the union in the scheme. Moved but still more than Parker: Some runners were in his classification tinamám closer to than others ratitům. Its "cohort Struthiones" Garrod is divided into four "families": one shared with ostrich rhea (Struthionidae), other cassowaries and emus (Casuaridae), third he himself kiwi (Apterygidae) and fourth were tinamy ( Tinamidae ).
Mivart (1877) has contributed to a better understanding of anatomy kiviho (see article by D. Naish the tetrapod Zoology ). Unlike Richard Owen, who is allied with tabonovitými , but it's only reinforced the fact that the kiwi fall between Revert. Mivart coincided with Huxley that the runners are a natural (monophyletic) group, and treated them as a single "family". Speculate about which birds ratitům should be the closest, however, offered the hypothesis that internal family relations. He presented it in the form of a tree diagram - which represents some progress from Huxley and anticipates that Fürbringera and his revolution in bird taxonomy. In the combined dendrogram Mivart ostrich (Struthio ) and rheas (Rhea ) casuar ( Casuarius ) and emu (Dromaius ) and finally kiviho ( Apteryx) and moa ( Dinornis ). The familial bond has defended Mr Owen, but to get Huxley's classification. Mivart (1877) further assumed that the line of kiwi-moa, cassowaries and emus are sister groups. Like Huxley and Mivart (1877) theorized that the runners never not fly and thus were primarily flightless. This view also evidenced in his second work of the same year (Mivart 1877b).
Sclater (1880) introduced a rather conservative classification that ignored the fossil and living birds divided into the usual "sub" Ratitae and Carinatae . Revert Sclater further divided into Struthiones (including ostrich and rhea), Casuarii (composed of cassowaries and emus) and Apteryges (kiwi). Somewhat more interesting was the system Stejnegera (1885), whose fundamental division of birds I write in connection with hesperornity below. Stejneger Huxley "superorder" Dromaeognathae divided into Struthiones (ostriches, rheas, [emu, cassowaries +] MOA) Aepyornithes (elephant birds of Madagascar), Apteryges (kiwi) and surprisingly Crypturi (tinamy). In Stejnegerově then distinguish the basis of what Pycraft 15 years later, naming Palaeognathae and what happens to a solid foundation all future phylogenetic hypotheses. Notice that in Stejnegerově of no group that would include all the runners, while excluding tinamy. Today's terminology, we could probably say that the author regarded as paraphyletic Revert to tinamám. Stejnegerovy exact opposite of anticipation, the system Reichenow (1882), which is not satisfied with the current division of birds into two or three "sub" and instead introduced seven so-called "series" that crowded living birds often bizarre uskupení. Pro běžce, úsporně seskupené do jedné "čeledi" Struthionidae, zavedl Reichenow synonymum Brevipennes.

3.5 Hesperornithes

    Ke konci 70. let 19. století se začínalo zdát, že mezi běžce brzy přibude nový - a extrémně zajímavý - přírůstek. Měl jím být Hesperornis , asi dva metry dlouhý, evidentně dravý pták s ozubenými čelistmi a silnýma nohama ze svrchní křídy dnešního Kansasu, který měl stejně jako běžci reduced wings and breastbone without the keel. (The degree of reduction of the wings could be classified somewhere between kiviho and birds Moa - his skapulakorakoid still meshed with the upper arm, but that it ended.) Hesperornis described by Marsh (1872), who also soon began to wonder where the development tree of birds this particular animal fit. Although often mentioned in literature suggest that Hesperornis and its relatives (example Hesperornithes or Hesperornithiformes) were close relatives of grebes (Podicipedidae), Loon (Gaviiformes), or groups made up of two previous positives (Gauthier & de Queiroz 2001; Hope 2002), somewhat oblivious to the fact that the earliest speculation about their close ratitům. Marsh even went so far as to Hesperornis called "carnivorous, ostrich floating" (Carnivorous, swimming sharp). If I understand the first time Marsh (1878), this term was used in an article for the magazine Popular Science Monthly , but eventually it also included in his detailed monograph Odontornithes (Marsh 1880), dedicated all fossil toothed birds of America .**

reconstructed skeleton of the carnivorous aquatic alleged Marsh ostrich Hesperornis regalis , issued by the Canadian Fossil Discovery Centre. (Source: wikimedia.org)

In it he notes Marsh different "pštrosovitých" characters hesperornisově on earth, and proposes two different explanations of them. The first traits that any indicate a relationship, and the runners hesperorniti is both inherited from a common "reptilian" ancestor - today's terminology, therefore, were pleziomorfie. Marsh describes, in this scenario would hurt hesperornitů ancestors looked like modern loons. During evolution, their wings reduced to the level of penguins, and then it was lost altogether, with the way their role in place of the swimming legs. The second explanation is really close relationship with the ostriches. The author writes:

Another Explanation He Seems The Whole Sea Reasonable, and more in accordance with the known facts. The Struthious characters, seen in Hesperornis , BE should probably regarded as evidence of a real affinity, and in this case Hesperornis Would Be a Carnivorous Essentially, swimming blades.

Marsh 1880:114

hesperornity with Marsh as a runner, but rather hard hit. Already Vetter (1885) supported his less popular alternative, according to which Hesperornis evolved from ancestors karinátních reduction of the wings and pectoral fin. If I know (I draw here ), Vetter speculate on hesperornisových closest relatives of this task, however, took an American ornithologist Stejneger (1885), the classification tends to be somewhat forbidding place a large number of new and completely unnecessary names, also criticized the time (Sharpe 1891). As if not enough already annoying enough "empty ranks (Rheiformes, Rheidae and Rhea have the exact same content), Stejneger even bothered with the fact that they at least have the same etymological basis: for archeopteryga retained both taxa Saururae Haeckel, but created it new, redundant "order" Ornithopappi. Leaving aside, however, these formal deficiencies Stejnegerův system is quite interesting. Birds now divided into four "sub", two of which he borrowed from the Marsh and the third from Haeckel. In his memoir Odontornithes divided Marsh (1880) toothed birds to people whose teeth grow from the groove (Odontolcae) and those with dental separate wells (Odontotormae). Stejnegerův system, this division follows:

"sub" Saururae
- "order" Ornithopappi ( Archaeopteryx ,? Laopteryx )
"sub" Odontotormae
- "order" Pteropappi ( Ichthyornis , Apatornis )
"sub" Odontoholcae [sic]
- "order" Dromaeopappi ( Hesperornis and relatives)
"sub" Eurhipidurae (wild birds)

Stejneger was perhaps not the first author who explicitly excluded hesperornity of logs made up of all living birds. This example is now called the name Neornithes (or Aves , sensu Gauthier & de Queiroz 2001), that it be done to Gadow in 1893 - Stejneger is perhaps the first to be explicitly named .*** This is undoubtedly right, in its time, but acceptance. It is interesting that with something like Stejnegerově system of four "sub" came across the Atlantic Menzbier (1887). He agreed with a special status and plazoocasých "odontotormů, wild birds, but tore into three" sub " - Revert to the historically unique, Karin and penguins (Eupodornithes).
With that in mind Marsh runner Hesperornis jurisdiction is erroneous, and further agreed (Fürbringer 1888; Thompson 1890). Fürbringer its gigantic monograph, of which more will be discussed, introduced to spur American water birds still use the name Hesperornithes (and Enaliornithes while Enaliornis today hesperornitem) and combined them with a purported asset named Colymbo-Podicipites, consisting of loons (Colymbidae today Gaviidae) and grebes (Podicipidae today Podicipedidae), the "suborder" Podicipitiformes. By contrast, Marsh side as one of the few built Newton (1884), who modeled Huxley categorized birds Saurarae taxa, Ratitae and Carinatae . Newton believed that the derivation karináti runners and lost teeth independently, while "plazoocasí (Saururae) they all possess. In accordance with the shared Revert to the carina and gear teeth. Under toothless runners, all the living representatives, birds MOA (Immanes) and elephant birds of Madagascar (Aepyornithes). Presumed toothed runner Newton (1884) divided into those with bikonkávními vertebrae and the vertebrae with saddle (which belonged to Hesperornis ), resulting classifications are as follows:

"sub" Saururae
- Archaeopteryx
"subclass" Ratitae
- teeth
  - with bikonkávními vertebrae (no known representatives)
  - the saddle vertebrae ( Hesperornis )
- without Teeth
  - Aepyornithes (= Aepyornis , elephant birds of Madagascar)
  - Apteryges (= Apterygidae, kiwi)
  - Immanes (Dinornithidae + = 'Palapterygidae ": moa birds, Palapteryx = Dinornis )
  - Megistanes ( = Casuariidae, emu, cassowaries +)
  - Rhea (= Rhea, rhea)
  - Struthiones (Struthio = , ostrich)
"sub" Carinatae
- teeth
  - with bikonkávními vertebrae ( Ichthyornis )
  - the saddle vertebrae (No known representatives)
- without teeth (living neognáti + tinamy)

reason for this arrangement, which today - just like any other system, allowing for the taxa hitherto undiscovered pre-defined properties - seems slightly strange words, the mismatch between what the former states of characters ornithologists and what to expect at Hesperornis and ichtyornise occurred in reality. As Newton (1884:36-7) explains, Ichthyornis had biconcave (in modern terminology amficélní) vertebrae and teeth located in the wells (such teeth is called tekodontní), while Hesperornis vertebrae had saddle and teeth set in a continuous groove. The problem is that it features the then regarded as secondary authors (teeth in the saddle-shaped vertebrae, and wells) did not occur together. Newton, although he admitted that the combination of archaic and derived characters is exactly what we expect evolutionary forces, but since both karinátů and Revert scored at least one pleziomorfický status symbol, he had to assume the convergent loss. With Newton's classification totally agree Pavlow (1885), which included Marsh Odontolcae taxa (= Hesperornithes ) between the runner and Odontotormae (= Ichthyornis ) to Carinatae .
Today we seem to Newton's view is flawed in many aspects. Not only that Hesperornis not to ratitům and Ichthyornis to karinátům, but both are outside the crowning example (= group of all living) birds, which Ichthyornis is closer to him (Clarke 2004). We know that all the ancestors of birds until the very distant past had tekodontní teeth (crocodiles and birds in their teeth wells inherited from the same ancestor) and that the present state of Hesperornis is thus derived, not primitive. Likewise amficélní vertebrae that have plagued the Marsh, although Ichthyornis neptačími shared with dinosaurs, but because of hesperornitů, patagopteryga, apsaravise and to some extent known konfuciusornitidů heterocélní vertebra ("saddle"), is more economical not interpret it as a common primitive character (sympleziomorfii), but derived (autapomorfický) reverzál, return to the original state (Clarke 2004). Since
hesperorniti around this time runners emerge from the story and never returned to him, yet leave you strictly followed the time line and jump to the "future" to show how the above-cited conclusions reached. I draw from Houde'a (1987), which views the development of competence hesperornitů briefly revised. Fürbringerova classification was the fate of more systematic hesperornitů considerable influence, but concreted it (as well as other views on avian evolution) and Heilmann (1926), who demonstrated the similarity of the red and hesperornitů rejected linking characters with runners. His position was widely followed (Lambrecht 1933, Brodkorb 1963). Already Storer (1960), but did not agree and Gingerich (1973, 1976) again drew attention to the runners hesperornitů features, namely their paleognátní floor (see previous article ). Unlike Marsh realized that is a primitive and not derived similarities, and argued that paleognátní type of floor is the primitive living birds, which led in his later article (Gingerich 1976) Cracraftovu questioned the hypothesis of monophyly of the runners. † Cracraft (1982) also came up with a bizarre twist, when using the wrong examined with cladistic analysis demonstrated the monophyly "gaviomorfů", consisting of hesperornitů, grebes and loons. Relationships with Grebe But at least in the case conclusively disproved already baptornise Martin & Tate (1976) and later examined with cladistic analysis carried out clearly both vykrývaly Hesperornis and ichtyornise among wild birds (usually in a trichotomy).

3.6 Primary and nelétavost polyfylie birds

Huxley came with two hypotheses, around which, in the years developed considerable discussion: 1) runners are monofyletičtí and 2) runners are a relic of period, the birds still could not fly, and Glider karináti their descendants. Marsh and Newton agreed with both statements, although Newton (1884) have already felt the need to defend him. The text also gives us a nice picture of how, when scientists thought:

It Has Been for Some Time and the Question Whether the ratite is degraded, and descended from the type of Carinae, or Carinae and the superior development of the ratite type. Eminent Several zoologists have Declared themselves in Favour of the Former Probability, and at first sight most people Would Be Inclined to DECIDE Them with [...]. But the easiest answer is not always the true one, and that the writer should present it before Seems That Be Answered this question and the reply given it Should Be Another-was the first Which animal Properly Could anyone call and "Bird," as distinguished from a "reptile," possessed of a keeled sternum or not? Now Would Birds seem to have been Differentiated from Reptiles [...]. There Is No Reason to Think That That period at [...] Any reptile snake and a keeled sternum. Hence it Seems Almost Impossible That The First Bird should have possessed one; That is to say, it must have been practically of the ratite type.

Newton 1884:43

Revert But if separated from the rest of the birds before this group had years, at what stage of evolution it actually was? Vogt (1879, 1880) to archeopteryga basis of his research - which could make for surprisingly little bird characters - suggested that runners have evolved from dinosaurs, while the karináti archeopteryga and one again in a kind of generalized reptile. (As noted Witmer [1991], this generalized reptile later authors in the interpretation of Vogt's hypothesis mistook the scaly, or lepidosaury.) This hypothesis now seems to be absurd - and the independent development of runners' karinátů "lizard like reptile from the required number of absolutely convergence inadequate number of differences that might explain. Polyfylie extreme hypothesis but support other authors, including Mivart (1881), in which the runners were descendants of dinosaurs and pterosaurs karináti again.

opposite view offered by Lindsay (1885), studying bird's breast bone. According to the keel karinátů relatively late phylogenetic structure, evolving along with powerful pectoral muscles serving to fly and since found remains of the keel of the baby rheas, concluded Revert that birds are descendants létavých.

Difyletičtí birds and their evolution under Wiedersheima. (Source: Witmer 1991: Figure 3, the digitized books.google.com)

Contact hypothesis popular in the 70 and 80 the 19th century into a single consistent model attempted wiedersheimi (1882, 1884, 1885, 1886). He said it was necessary to seek a common ancestor with other birds Revert to Early Triassic, or even the late Palaeozoic (!), While the front was also the ancestor of dinosaurs and pterosaurs. Wiedersheimi to be featured again as an indeterminate reptile (which was again later misinterpreted the "lacertiliana, see eg de Beer 1954), which gave rise to one of the scaly animals, which are split with the pterosaurs and the line leading to birds. On it lies Archaeopteryx, from which evolved all later birds including ichtyornise, which in turn is the ancestor of all living karinátů. On the other hand, Wiedersheimova of "deep ancestor" evolved dinosaurs, whose descendants include Hesperornis and Revert. It is not hard to see that this strange hypothesis was based solely on the authority of other scientists, like the Marsh and Vogt.
The loss of popularity suffered a bird difylii hypothesis is most likely responsible Fürbringer. In 1888 he published his monumental work Untersuchungen zur Morphologie und Systematic der Vogel , in two volumes with a total of 1751 Parties presented a phylogeny of birds not only through hierarchical lists (such as those mentioned above), but also the development tree. The scope and quality of this work also suggests that - as recently pointed Mayr (2011) - neither Livezey & Zusi (2007) in his numerical analysis of 2954 morphological characters did not get the cladograms much different from that which Fürbringer presented 120 years ago . Fürbringer also presented compelling evidence to suggest that all birds - including the Revert - Glider and were initially speculated that bird flight originated in trees (arboreal hypotéza).

3.7 Od Fürbringera po Gadowa

    Co se týče systematiky běžců, Fürbringer (1888) pokládal tuto skupinu za parafyletickou - dosud přitom soupeřila Owenova hypotéza o polyfylii s Huxleyho hypotézou o monofylii. Pštrosi ( Struthio ) mají na jeho stromku ke zbylým ptákům nejdál - hned po archeopterygovi, který je nejbazálnějším nehypotetickým ptákem ve Fürbringerově stromku. Následuje taxon "Hippalectryornithes" (autor měl opět onen nepříjemný zvyk používat vícero redundant names for one and the same example), including cassowaries, emus and dromornitidy - large birds of the Australian Oligocene to Pleistocene, which, while reminiscent of the runners, but today are considered vrubozubé. Even today they are closer relatives to birds létavým nanduové ( Rhea). Finally, the kiwi (Apteryx ) is falling along with the MOA (dinornitidy) in the stocks named Apteryges, whose position relative to other birds somewhat obscures the artistically impressive but a bit impractical to implement tree. Fürbringer attempted a synthesis of current hypotheses on the phylogenetic position of Tina and her tree is therefore set in (under the name Crypturi) to polytomie with hrabavými and Apteryges. Gussekloo (2000) states that Fürbringera impressed by the morphological diversity in bony běžčím floor, Huxley by his notion of "dromeognátní type is not fully appreciated.

Development Tree birds according Fürbringera, which is evident in both runners parafylie (different to the rest of the runners have different birds away), both of secondary nelétavost (Glider Archaeopteryx is located below the tree). On the right, then the clearer version of the same tree, reprinted Mayr (2011), applying the current nomenclature and omitting fossil taxa. (Modified from Sharp 1891: Plate IV [digitalized archive.org] and Mayr 2011: Figure 2)

The Fürbringera Gadow continued, addressing it from the 70 of the 19 century form of bird intestinal loops. The only consensus in this character set by Gadow (1889) for Revert discovered the second clockwise loop and left-handed third. These characters, however, also show tinamy, Galliformes, and hoacin crested cuckoo. Apparently the findings thus confirm Fürbringerovy, but as noted by Sibley & Ahlquist (1972), despite the fact that it was by far the biggest promoter of Gadow Fürbringerovy work in the Anglophone world, in many respects, he disagreed with it - Revert and classification was one of them.

Seebohm (1890) has since presented the work kiviho which came back between runners (in his terminology strutioniformy) and rejected their relationship with hrabavými tinamami and, as suggested Fürbringer. Seebohmova work in the coming years, underwent considerable criticism, especially from Newton. As a "reactionary" but rejected it as Gadow (1893), when he published his groundbreaking work systematically. In it applies the principles of a primitive kind of numerical analysis, including the weighting of characters, it explores many different aspects of bird anatomy and bird among other things, puts up the Crown Example name Neornithes. *** Despite his previous findings on the construction of the intestines are monophyletic Gadow recognizes Revert. From Fürbringerova positives ( Tinamidae , Galliformes, Apteryx ) Gadow kiviho broke and sent him back to the runner. These steps should be used for further taxonomy of birds of paramount importance: Gadowova classification will be subject to minor modifications (Beddarda, Wetmore'a and others) used so long as cladistics and molecular phylogenetics not make unnecessary classification.

* I know the date it attaches some importance exaggerated, but personally I do not have much to consider it as the beginning of modern zoology as a whole.

** Just in passing: that deletions of the budget for science and research - especially a science and research that have extensive practical applications (eg, studies of fossil birds) - have a long tradition in history, evidenced by the fact that a deputy the state of Alabama Hilary Herbert in 1892 used a 44-page excerpt from Marsh monographs directly on the soil of the House of Representatives as the ultimate example of throwing taxpayers' money. He even passed around a luxury edition Odontornithes , printed on Marsh's own expense, as evidence of profligacy U.S. Geological Survey.

*** Or not. Search priority designation can be finished suffering, as in the earlier literature is simply not clear what the authors intended the asset's name together. After some time as covering all living birds - and the only wild birds - taxon Ornithurae , Haeckel already erected in 1866. Today everyone will agree that this includes several taxa outside the crown of literature is but a name used for three different strengths: Chiappeho node-based ( Hesperornis + wild birds), Serenův branch-based (all more than sparrows archeopterygovi) and Gauthierův and de Queirozův apomorphy-based (all birds with short tails bearing bone pygostyl, homologous with that of the Condor). Why? Since it is not clear how this name konceptualizoval Haeckel. A similar case Even the name is Neornithes . It is usually accepted that Gadow (1893), it intended to explicitly name the group composed of all living birds, but Gauthier & de Queiroz (2001) argue that it was going to pay for Haeckelovo Ornithurae , and suggest that the extended hesperornity ichtyornise and since there Gadow included (considering that it is a crown I lay members).

† It is fitting to note that Houde (1987), however, disputed that, as described by Gingerich Hesperornis is true paleognátní patro. 

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