|
"Most scientific breakthroughs are nothing else
than the discovery of the obvious", Eberhard Zangger comments his well-founded
opinion that Atlantis is merely a saga about the Trojan War, distorted by
translation errors into Egyptian and back. I leave it to you to judge whether my
paper is a breakthrough in science or in blockheadedness, but before I add a
disclaimer: I haven't seen any of the fossils I'm writing of, only the
Confuciusornis specimen at the Natural History Museum here in Vienna
(and it didn't support or falsify any of my opinions; I couldn't look very
close), I have derived everything from the literature. Therefore everyone else
could have arrived at my conclusions, at least some of which are pretty obvious
to me, by simply reading all the papers and websites (dinosauria.com,
Dinosauricon, archives of this list) I have used. Indeed, a few have been
reached by others (remember Details on Microraptor where HP Mickey
Mortimer found Archaeopteryx closer to dromaeosaurs than to birds?) My
ideas are few, most of them is putting the published pieces
together.
Parts of the paper are even outdated: The Early
Jurassic dromaeosaur teeth from Antarctica, Microraptor,
Yandangornis (for which I hardly had information) and Avimimus
(which I thought to be a chimera) lack.
I have copied much of the paper, because I can best
paraphrase my own words in my own words...
I still can't give a complete citation, as I
haven't received the Dinosaur Society Quarterly Journal 4.1 yet. Has anyone
else found it?
The abstract is very short (because the paper is
very long):
"Archaeopteryx completely lacks bird
synapomorphies; therefore the cladogram of derived coelurosaurs is rearranged to
(((Metornithes + Oviraptorosauria) + Arctometatarsalia) + (Archaeopterygidae +
Dromaeosauridae))."
I should have inserted a few words on the origin of
flight...
Archaeopterygidae is to mean A.
and Unenlagia ?= Megaraptor ?= Unquillosaurus (possible
synonymies after the Dinosauricon); analyses by HP Mickey Mortimer put
U., at least, in the Dromaeosaurinae.
The main paper begins with "When
Archaeopteryx was discovered, birds were thought to have lots of
autapomorphies, many of which were recognised in Archaeopteryx. But for
the past 20 years or so, former bird autapomorphies have been 'sliding down the
cladogram', so it has now become very difficult to diagnose a
bird."
Explicitely (most of this not new to anyone on this
list, I think; I hope this lines up correctly on your screens):
Former bird autapomorphy found in A. |
Argument that it has slided down the cladogram
opisthopubic
pelvis |
is orthopubic in A. and Patagonykus, opisthopubic in
dromaeosaurs [and secondarily
orthopubic in
U., if HP Mickey Mortimer is right) => probably prone to
convergence
furcula |
(you all know)
feathers
| (you all know)
wing and tail feathers
| (you all know)
unserrated teeth with
constriction | you know,
Pelicanimimus, Byronosaurus... "Elzanowski & Wellnhofer
(1996) note that the
between root and
crown teeth
of Archaeopteryx itself only sometimes have a slight constriction, and
no sign of an
expanded
root like that occuring in Ichthyornis. They seem to be an adaptation
for
eating
fish, not a feature useful in cladistics."
Former bird autapomorphy found in A. |
Argument that it has migrated up the cladogram
loss of the postorbitals
|
A. has postorbitals, only euornithines, it seems, don't
various forms of skull
kinetics |
all unknown in A., because no complete A. skull is
known;
seems rather improbable from what is known
loss of coronoid (III)
| "In fact, this feature has a very spotty
distribution – Archaeopteryx and
ornithomimids
lack a distinct coronoid, Oviraptor has a tiny one, while
Dromaeosaurus
and Velociraptor (Currie, 1995) still have two (fused
coronoids
II and III), which is considered highly plesiomorphic."
Blue emphases, inserted for easier
reading, not in the original.
Brackets [] are in the original, for commenting I
use <> here.
"In addition, it has been discovered that
Archaeopteryx lacks some features that birds share with
arctometatarsalians and oviraptorosaurs. Sometime it is mentioned that the
latter three taxa (troodontids and tyrannosaurs: Currie, 1995, oviraptorosaurs:
Sues, 1997 contra Norell et al. 2000) have a separate exit
for the cranial nerve V1, which normally exits the braincase through a
large foramen together with V2 and V3. Dromaeosaurus (Currie, 1995) and
Archaeopteryx (Elzanowski & Wellnhofer, 1996) exhibit the primitive
condition. Arctometatarsalians and oviraptorosaurs also share pneumatised quadrates and articulars not occuring in
Archaeopteryx (Elzanowski & Wellnhofer, 1996) and
Dromaeosaurus (Currie, 1995). Andrzej Elzanowski is sometimes referred
to as having discovered that some features of the
oviraptorosaur palate are more birdlike than that of
Archaeopteryx. <Has been published in detail in the meantime, I
still haven't found the paper.> Osmólska & Maryanska (1997) note that
oviraptorid quadrates are double-headed, i. e. they have an additional contact to the
braincase, and they are much more pneumatised than in
arctometatarsalians (Osmólska & Maryanska, 1997). <I have posted
the relevant quotes a few weeks ago.> Both are features occuring in birds but
not in Archaeopteryx (Elzanowski & Wellnhofer, 1996, Chatterjee,
1997, Paul 1997) <...>. Oviraptorosaurs (Sues, 1997) (including
Caudipteryx <and Protarchaeopteryx?>) also share with
alvarezsaurids (Novas, 1997) rather short tails that are not
stiffened distally, a condition seen in no other theropods. <...>
Pneumatic features also fit this picture: Archaeopteryx is no more
extensively pneumatised than dromaeosaurids, i. e. cervical and dorsal vertebrae
as well as their ribs are pneumatised (Britt et al., 1998). In oviraptorosaurs
(Currie, 1995) and alvarezsaurids (Novas, 1997), pneumatic
foramina (or depressions) occur as far back as the middle of the tail!
(In recent turkeys (Britt et al., 1998), the sacral and free caudal vertebrae
are pneumatised, in ostriches (Britt et al., 1998) the caudals are not.) With
the description of Nomingia (Keesey, 2000, /genera/nomingia.html),
the pygostyle has joined the list of maniraptoran
(see cladogram) synapomorphies (no alvarezsaurid tail end is known,
Caudipteryx and Protarchaeopteryx could have pygostyles
<just discussed>, judging from photographs (Ackerman, 1998), the
resolution of which is too coarse to be certain)."
"A commonly
cited bird apomorphy in Archaeopteryx is the reverted hallux; all preserved feet are crushed, the joint
surface between metatarsals I and II is destroyed [in e. g. the London specimen
and not prepared in the Berlin one], thus to what extent the halluces were
reverted is still a matter of debate. In any case, Archaeopteryx was
incapable of perching, because the halluces were too short and situated too high
on the foot. <Both characters not present in Microraptor... only
we don't know to what extent its halluces were reverted
either.> Caudipteryx dongi (Normile, 2000), on the other hand,
seems to have a reverted hallux as long as the other toes <I have meanwhile
found, but not yet copied, the original Vertebrata PalAsiatica paper; it wasn't
that long> – its description will fuel lots of additional debates <for
which I am waiting>. A single bird synapomorphy, according to Elzanowski
& Wellnhofer (1996), has been left to Archaeopteryx: the triradiate palatines. I think that it is most parsimonious
to explain this feature by convergence <ô surprise>, though I
have no idea why this should have occurred."
Birds down to Alvarezsauridae have prokinesis
(Chiappe, Norell & Clark, 1998); confuciusornithids have akinetic skulls
(Hou et al., 1999), which is probably secondary because the jugals don’t have
ascending processes.
The cladogram includes all Ornithodira, so most of
it is simply copied from elsewhere. Let's see whether I can write it
Dinosauricon-style:
--Ornithodira
|---Pterosauromorpha
`--Dinosauromorpha
|--Lagerpeton
`--Dinosauriformes
|--Lagosuchidae
`--+--Pseudolagosuchus
`--Dinosauria
|--Ornithischia
`--Saurischia
|--Sauropodomorpha
`--Theropoda
|--Herrerasauridae
`--Neotheropoda
|--Ceratosauria
`--Tetanurae
|--Spinosauridae
|--Torvo-/Megalosauridae
`--Neotetanurae
|--Carnosauria
`--Coelurosauria
|--Dryptosaurus, Diplotomodon, Deltadromaeus?
|--Compsognathidae
`--Eumaniraptora
|--suggestion:
Archaeopterygiformes
| |--Archaeopterygidae
(see above)
|
`--Dromaeosauridae
`--Maniraptoriformes
|--Arctometatarsalia
|
|--Tyrannosauroidea
|
`--Bullatosauria
| |--Troodontidae
| `--Ornithomimidae
`--Maniraptora
|--Oviraptorosauria
`--Metornithes
|--Alvarezsauridae
`--Pygostylia
"Cladistic implications
Archaeopteryx has no bird synapomorphies,
and is probably nothing more than a glorified small flying near-dromaeosaur,
'[a]nd in spite of the fact that dromaeosaurids are often proclaimed to be the
most birdlike of the theropods <...>, they lack many of the theropod-avian
synapomorphies found in other theropod families, and have too many
specialisations to be plausible avian ancestors' (Currie, 1995, page 587).
Considering this along with the 'theropod-avian synapomorphies' listed above
<below> produces a cladogram <...> which differs from all
others in the following features: oviraptorosaurs are the closest relatives of
birds, with which they form the taxon Maniraptora (conventionally defined as
Neornithes > Ornithomimus, in Sereno [1998] defined as
Oviraptor + Neornithes; at present, these definitions do not change the
contents of Maniraptora). Maniraptora and Arctometatarsalia form
Maniraptoriformes, which has been defined as Ornithomimus + Neornithes.
The taxon with the definition Deinonychus + Neornithes has recently
been named Eumaniraptora. Its name implies that it was meant to be a part of
Maniraptora, whereas now it is vice versa -- the irony of cladistic
definitions.
I propose to name the taxon which includes
Archaeopteryx and dromaeosaurids Archaeopterygiformes, which is among
the oldest available names, and to define it as Archaeopteryx >
Neornithes. Sereno (1998) has defined Archaeopteryx this way – that is
why I consider it useless to define genera.
But what about birds themselves? A commonly used
definition is Aves: Archaeopteryx + Neornithes (e. g. Sereno 1998) and
Avialae: Neornithes > Velociraptor. In this case, Aves is <in
terms of content> a senior synonym of Eumaniraptora and Avialae one of
Paraves ( = Neornithes > Oviraptor [Sereno, 1998]), and, whether
Aves or Avialae is called 'birds', Tyrannosaurus is a bird. Now the
question is whether we accept animals lime Tyrannosaurus or
Ornithomimus as birds or whether we allow the
definitions to change <we can do the latter when PhyloCode is enacted>. In
the latter case I propose to define Avialae as Neornithes >
Oviraptor, Ornithomimus, Velociraptor and other
well-known non-avian theropods and to forget Aves until an animal is found
(perhaps Rahonavis or Protoavis, when they are <will be, I
mean> be better known and studied -- see below) being closer to Metornithes
than to Oviraptor. Metornithes is unaffected by the change of the
position of Archaeopteryx and is a useful taxon that continues to be
defined as Mononykus + Neornithes."
The following part is largely copied from the
version on my computer where I wrote it; I have added a few slight changes,
which I can't reconstruct because I would have to find the old emails which were
made nearly inaccessible in the last few computer crashes, and the editor, Jeff
J. Liston, has corrected a few mistakes in my English. So the actual published
paper is minimally different.
"Evolutionary and palaeoecological
implications
If it is true that Archaeopteryx, being capable of
flight, is less closely related to birds than many flightless genera, then all
the current theories ( = not much more than the cursorial and the arboreal ones)
on the evolution of bird flight and feathers are in serious trouble, and even
Baron Nopcsa's hypothesis that wings increase running speed, which was repeated
as late as 1999 (Burgers & Chiappe, 1999), cannot save the former <but
that paper can possibly save A.'s ability to take off from the
ground>. But, as Ebel has shown in 1996, they are in trouble anyway, and
there is already an alternative (Ebel, 1996):
First,
protofeathers evolved as a protein sink: Bird ancestors, early tetanurans,
theropods, dinosaurs or ornithodirans in general are commonly thought to have
been insectivorous. Insects, which radiated in the early Mesozoic, contain much
more fat and protein than vertebrates and thus are a good reason for becoming
and staying endothermic. (Let’s simplify the case and assume that all
ornithodirans were endothermic – Paul & Leahy [1994] have shown excellently
that even the largest ones were forced to be.) But insect protein contains more
sulphur than small endotherms can use or excrete. So these animals deposited the
sulphur-rich amino acids in their scales, which are made of keratin, a protein
containing up to 3 % sulphur. The scales were enlarged and split during growth,
used for insulation and display, and sometimes they were shed to get rid of the
sulphur. This is a wonderful explanation for why birds moult (Reichholf 1997
<and others, which may be cited in the published version>).
Sinosauropteryx, Sinornithosaurus and, if you kindly forget its diet,
Beipiaosaurus can be considered examples for such animals. Because the
scutes on bird feet are often considered to be modified feathers, it is possible
(Poling, 1996) that feathers are in fact an ornithodiran synapomorphy, that
pterosaur hairs are protofeathers and that the typical polygonal scutes of many
skin impressions from large dinosaurs show that large dinosaurs lost feathers
several times independently like large mammals which lost hair. Polar
hypsilophodontids from the Early Cretaceous of Victoria/Australia are often
illustrated bearing feathers which they could surely need.
After that,
some tetanuran invented lying on its eggs and holding them with its arms. The
arms of the famous brooding Oviraptor and ?Ingenia skeletons all
circle the eggs in a way that if they had borne wing feathers, they had covered
the nests and shielded them from sun and rain (Hecht, 1998 <yes, HP Jeff
Hecht who mentioned this hypothesis by HP Tom Hopp in New Scientist>).
Apparently there was a strictly Darwinian advantage in lengthening the arms and
the feathers: longer arms = more wing area = bigger nests = room for more eggs =
more offspring."
May be a good argument, at least it sounds like
one, doesn't it? =8-)
"Then one of them, maybe the first eumaniraptoran,
started eating fish instead of insects. For catching fish it is useful to be
able to swim. <Ô surprise.> How would a tetanuran have swum?
Undulating the tail, which many have suggested, was impossible – the distal half
was stiffened – and would have been ineffective, because dinosaur tails were
always pointed at the end instead of flattened like in crocodiles. Using their
legs would have been ineffective, too, because they weren’t able to sprawl and
their toes were not webbed. There was only one other possibility – using the
arms, which already bore wing feathers . Thus this species learned to fly
underwater like the dippers, which have only one adaptation to their lifestyle,
the ability to close their nostrils with skin that cannot fossilize. <In
Ebel's paper wing feathers evolved for swimming -- rather improbable IMHO,
but *asymmetrical* wing feathers probably evolved at this stage.> Flight
capabilities were able to be steadily sophisticated without the danger of
falling down, in contrast to flying in air, which must be nearly perfect from
the beginning. At the same time, the tail was lightened, stiffened further and
got long, stiff tail feathers for steering (Ebel, 1996). The semilunate carpal
may also have evolved at this stage.
If a
large-winged animal can fly underwater, it can also fly in air, because in air
there is much less drag. Ebel (1996) has calculated that flying at a speed of
0,35 m/s underwater produces the same dynamic pressure as flying at 10 m/s in
air. Thus, after having got a fish, it’s easier to reach the shore by flying in
the air than underwater. Being able to fly is also an advantage in such a
lifestyle because "the range of accessible fishing grounds […] [can be]
enlarged" (Ebel, 1996, page 279). This condition is beautifully seen in
Archaeopteryx. Note that Archaeopteryx was unable to glide: its
long tail positioned the centre of gravity in the hips, which is useful for
bipedal running and unimportant in underwater flying, but requires constant
flapping in air. Ornithothoracines have reduced the tail to place the centre of
gravity between the shoulders, at the same place as the centre of lift, and
therefore can glide and soar. (Ebel, 1996)
Dromaeosaurids became secondarily "flightless". <We know this from somewhere,
don't we?> They used their tails for balancing, their hands for grasping
only, enlarged the claws on their hyperextendable second toes and changed their
diet. This seems to have been an instant success, since there is a possible
dromaeosaurid from the US Late Jurassic as well as some dromaeosaurid remains
from the UK Middle Jurassic (Ebel, 1996)."
As well as
other Middle and even Early Jurassic ?dromaeosaurs mentioned onlist by HP Mickey
Mortimer.
"Arctometatarsalians did the same, but they changed to long-distance pursuing,
as indicated by their feet. They were already present at the same time as
dromaeosaurids – there are the troodontid Koparion, some ornithomimid
teeth <and that Kimmeridgian finger from Great Britain -- thanks to the list
I mention it in the published version> and the probable tyrannosaurid
Stokesosaurus (Chure & Madsen, 1998) from the Late Jurassic of the
USA, and if Stokesosaurus is a tyrannosaurid, then Iliosuchus from
the Middle Jurassic of Great Britain, where troodontid teeth have been found
(Keesey, 1999, Poling [ed.], 1997 – 1999), is one, too (Keesey, 1999, Poling,
/clado/tyrannosauroidea.html <see below>). Therefore, eumaniraptorans,
including maniraptorans, must have diversified at that time or even earlier.
<...> Early Jurassic and even Late Triassic bird tracks are appearing all
over the world (e. g. Chatterjee, 1997, Gierlinski, 1996 <another paper, not
the 'feather' impressions>). Surely somewhere in the reader’s head there is
lurking the idea that Protoavis might fit this picture <...>.
Protoavis is a problem because of its poor preservation, and it is
probably a chimera, but it is very difficult to explain which part comes from
which animal. For instance, the "hand" looks much like an herrerasaur foot – but
only the first three digits: "metacarpal IV" bears no phalanges, and no similar
bone occurs in any known herrerasaur, as is the case with the "quill knobs" on
"metacarpals II and III". There is no explanation for the (though unfused)
furcula and the keeled sternum. Additionally, the chimera argument doesn’t
really work – if only a single bone attributed to Protoavis is avian,
then there was a Triassic bird (Chatterjee, 1997). The braincase at least is
surely tetanurine (Currie, 1995)."
Protoavis quite clearly has a common exit for the 3 branches of cranial
nerve V. Bad for me or bad for that braincase?
"Oviraptorosaurs must also have originated at that time, but the fossil record
keeps silent. <Not entirely true, see below.> They seem to have
changed their diet from fish to freshwater snails, which were abundant in
Mongolia and Liáoníng/China, and (like many birds later) from underwater flying
to wading (Chirostenotes is often said to have had wading adaptations). A
diet of mainly freshwater snails is probably the best explanation for their odd
skulls as well for the protruding premaxillary teeth of Caudipteryx which
might have been used in pulling snails out of mud.
Rahonavis (Forster
et al., 1998) states a problem for my ideas about theropod phylogeny. Its tail
looks just like that of Archaeopteryx, but the scapula-coracoid joint was
mobile, indicating that Rahonavis should be closer to ornithothoracines
and confuciusornithids than alvarezsaurids. The quill knobs are so small that
I’m sure no one has really looked for them in Archaeopteryx, so they
might be a general eumaniraptoran feature. Someone should find the skull and a
hand (where most relevant apomorphies are located), without which I cannot place
Rahonavis accurately."
Still
true. :-( Is it likely that a mobile scapula-coracoid joint can evolve twice (or
be a misinterpretation)? In this case Rahonavis could belong to
Archaeopterygiformes... MORE FOSSILS!!!
Yandangornis and Avimimus are probably maniraptorans because
of their similarities to oviraptorosaurs and alvarezsaurids.
"Note
added
The 7th specimen of
Archaeopteryx was made the holotype of the new species A. bavarica
based only on its ossified and fused sternum indicating maturity in contrast to
the other specimens in which the sterna were not ossified (the mysterious
8th specimen has not been described yet). I think the bone reported
as the sternum is just one (the left or the right one) and should
be rotated by 90°, which would explain its odd shape (for a theropod) and small
size (the gastralia don’t reach it according to [Poling [ed.], 1998,
/lungs.html]). Therefore, the 7th specimen may turn out to be
immature (it is a ?neotheropod synapomorphy that the sterna fuse in adults) and
the separate species A. bavarica to be unnecessary."
Has been contested onlist.
Guide to the cladogram
(shortened)
"Ornithodira: endothermy, insectivory,
parasagittal limb posture, bipedality, ?jumping locomotion, hairlike
feathers?
Dinosauriformes: running locomotion
Neotheropoda or earlier: furcula? (not ossified – like the sternum – in e. g.
compsognathids, attached to the scapula by ligaments and therefore frequently
falling off before burial, which explains its rare preservation [Makovicky &
Currie, 1998]; the ventral ends of the clavicles of Sinornithoides and
probably Carnotaurus are broken – perhaps the middle part of a furcula
has broken away [Mortimer, 2000 <in the List archives, can't find it now; it
is properly cited in the published version>])
Neotetanurae: fused distal carpal 1 + 2 semilunate to permit the swivel wrist
motion, 'terrible claws' ?convergently or ?primitively in Ornitholestes,
Archaeopterygiformes (the 2nd toes of Archaeopteryx are
hyperextendable, too [Paul, 1996]), Troodontidae, Confuciusornithidae (pers.
obs. and Ji, Chiappe & Ji, 1999), recent seriema (Cariama cristata)
and Rahonavis, the relationships of which are unknown beyond
Eumaniraptora (Forster et al., 1998) – see below <above>. Sereno (1998)
defends his usage of Neotetanurae by stressing the point that Avetheropoda was
originally <PDW etc.> used not as a node-based taxon
Allosaurus + Neornithes, but as a more inclusive stem-based
one.
Carnosauria: nearly triangular antorbital fenestra, some features of hand, lower
jaw, skull and pelvis according to Paul (1998); includes Allosauridae,
Ornitholestidae, probably Sinraptoridae and Carcharodontosauridae. <Any
comments???>
Eumaniraptora: nearly all cervical and dorsal vertebrae pneumatised to varying
degrees, furcula boomerang-shaped, shoulder joint facing laterally, arms
(theoretically) reaching at least the level of the ankles when fully extended
downwards, semilunate carpal enlarged, metacarpal I reduced in length and
metacarpal II long and slender to improve the opposability of the thumb, and
probably wing and tail feathers.
Maniraptoriformes: separate exit for the cranial nerve V1, articulars
and quadrates pneumatized.
Maniraptora: tail short (no more than about 30 vertebrae) and not stiffened
distally, quadrate extensive-ly pneumatised and double-headed, cervical, dorsal,
sacral, and proximal caudal vertebrae pneumatised, ornithoid eggshell
(Varricchio et al., 1997)
Archaeopterygidae (newly defined as Archaeopteryx >
Deinonychus) <may be redundant again>
Oviraptorosauria: loss of maxillary and dentary teeth, skull boxlike and adapted
to crush ?freshwater snails;includes Oviraptoridae, Caenagnathidae,
Microvenator, Caudipteryx and probably Protarchaeopteryx
(still not completely prepared). Thecospondylus with its ?7 or more
sacrals from the Isle of Wight and a femur from the same site (Naish, 2000) also
are probably oviraptorosaurian. Keesey (2000, /taxa/coelurosauria.html) mentions
an oviraptorosaurian caudal vertebra from the US late Jurassic.
Metornithes: loss of jugular ascending process, carpometacarpus
'Dryptosaurus etc.' = Dryptosaurus + Deltadromeus.
Holtz (1996) refers to this group as Dryptosauridae, but that term has already
been used for Dryptosaurus + Diplotomodon. Dryptosauria is
suggested as and alternative name for the group.
'Unenlagia etc.' = Unenlagia, ?Megaraptor,
?Unquillosaurus. Megaraptor and Unquillosaurus may be
junior synonyms of Unenlagia (Keesey, 1999).
Kakuru, a
tibia and a toe phalanx from the Early Cretaceous of Australia, could be an
alvarezsaurid, because it resembles the respective parts of Avimimus
which are thought to be mononykine (if Avimimus is indeed a chimera
<obviously not>). A probable Early Cretaceous caenagnathid is known from
Victoria/Australia, and there is a possible oviraptorid sacrum and ilium from
the same time of Brazil (33), which I guess to be possibly alvarezsaurid (for
biogeographical reasons)."
"Conclusions
The answer to the title question is 'No –
otherwise I wouldn’t ask.' Archaeopteryx lacks any bird synapomorphies as
well as synapomorphies shared by arctometatarsalians, oviraptorosaurs and birds.
Oviraptorosauria and Metornithes form (by definition) Maniraptora, which is
diagnosed by short tails lacking a transition point, backbones pneumatized back
to the middle of the tail, and extensively pneumatized double-headed quadrates.
Maniraptora and Arctometatarsalia form Maniraptoriformes, whose synapomorphies
include an extra exit for the cranial nerve V1 and pneumatized
quadrates and articulars. Archaeopterygidae and Dromaeosauridae are confirmed as
closest relatives; the taxon containing both is suggested to be called
Archaeopterygiformes. Archaeopterygiformes and Maniraptoriformes form (by
definition) Eumaniraptora, to whose synapomorphies can be added boomerang-shaped
furculae (reversed in Dromaeosauridae sensu stricto) and probably wing and tail
feathers. The relationships of Rahonavis and Protoavis remain
unclear. <...> – The separate species Archaeopteryx bavarica for
the 7th specimen is probably unnecessary.
References
<Have become more now and are published
in different layout.>
J. Ackerman: Dinosaurs Take Wing,
National Geographic July 1998, 74 – 99
B. B. Britt, P. J. Makovicky, J. Gauthier
& N. Bonde: Postcranial pneumatization in
Archaeopteryx, Nature 395, 374 – 376 (24
September 1998)
P. Burgers & L. M. Chiappe: The wing
of Archaeopteryx as a primary thrust generator, Nature 399, 60 – 62
(6 May 1999) S. Chatterjee: The Rise of Birds. 225 Million Years
of Evolution, Johns Hopkins University Press 1997Chen P., Dong Z. & Zhen
S.: An exceptionally well-preserved theropod dinosaur from the Yixian
formation of China, Nature 391, 147 – 152 (8 January 1998)L. M. Chiappe, M. A. Norell & J. M.
Clark: The skull of a relative of the stem-group bird
Mononykus, Nature 392, 275 – 278 (19 March
1998) D. J. Chure & J. H. Madsen: On the presence of furculae
in some non-maniraptoran theropods, JVP 16(3), 537 – 577 (September
1996)D. J. Chure & J. H. Madsen: An
unusual braincase (?Stokesosaurus clevelandi) from the Cleveland-Lloyd
Dinosaur Quarry, Utah (Morrison Formation; Late Jurassic), JVP 18(1), 115 –
125 (March 1998) P. J. Currie: New information on the anatomy and
relationships of Dromaeosaurus albertensis (Dinosauria: Theropoda),
JVP 15(3), 576 – 591 (September 1995)K. Ebel: On the origin of flight in
Archaeopteryx and in pterosaurs, same magazine as (21) 202(3), 269 –
285 (December 1996)
A. Elźanowski & P. Wellnhofer:
Cranial morphology of
Archaeopteryx: Evidence from the seventh skeleton, JVP 16(1), 81 – 94 (March 1996) A. Feduccia:
The Origin and Evolution of Birds, Yale University Press
1996C. A. Forster, S. D. Sampson, L. M. Chiappe
& D. M. Krause: The Theropod Ancestry of Birds: New Evidence from the
Late Cretaceous of Madagascar, Science 279, 1915 – 1919 (20 March
1998)
E. Frey & D. M. Martill: A possible oviraptorosaurid
[sic] theropod from the Santana formation (Lower Cretaceous, ?Albian) of
Brazil, N. Jb. Geol. Paläont. Monatshefte 1995 [which month?], 397 – 412
(I’ve only read the abstract in Zentralblatt für Geologie und Paläontologie:
Teil II – Paläontologie 5/6 [1997], 367.)
G. Gierliński: Avialian theropod tracks from
the Early Jurassic strata of Poland, Zubia 14, 79 – 87 (1996) (I’ve only
read the abstract in Zentralblatt für Geologie und Paläontologie: Teil II –
Paläontologie 5/6 [1997], 337 <so far>.)
J. Hecht: Let me take you under my
wing…, New Scientist (25 April, 1998), 22 Hou L., L. D. Martin,
Zhou Z., A. Feduccia & Zhang F.: A diapsid skull in a new species of the
primitive bird Confuciusornis, 399, 679 – 682
(17 June 1999)Ji Q., L. M. Chiappe & Ji S.: A new
Late Mesozoic confuciusornithid bird from China, JVP 19(1), 1 – 7 (March
1999) T. M. Keesey: The Dinosauricon, dinosaur.umbc.edu = www.dinosauricon.com = dinosauricon.comP. J. Makovicky & P. J. Currie: The
presence of a furcula in tyrannosaurid theropods, and its phylogenetic and
functional implications, JVP 18(1), 143 – 149 (March 1998)
D. Naish: A small, unusual theropod
(Dinosauria) femur from the Wealden Group (Lower Cretaceous) of the Isle of
Wight, England, N. Jb. Geol. Paläont. Monatshefte 2000 (4), 217 – 234 (April
2000) M. A. Norell, P. Makovicky & J. M. Clark: A
Velociraptor wishbone, Nature 389, 447 (10 October,
1997)M. A. Norell, P. J. Makovicky & J. M.
Clark: A new troodontid theropod from Ukhaa Tolgod, Mongolia, JVP 20(1),
7 – 11 (March 2000)
D. Normile: New Feathered Dino Firms Up
Bird Links, Science 288, 1721 (9 June 2000) F. E. Novas:
Anatomy of Patagonykus puertai (Theropoda, Avialae, Alvarezsauridae),
from the Late Cretaceous of Patagonia, Journal of Vertebrate Paleontology
17(1), 137 – 166 (March 1997)H. Osmólska & T. Maryańska: Mongolian Oviraptorids,
The Dinosaur Report Spring 1997, 1, 8 & 9
G. S. Paul: Predatory Dinosaurs of the
World, Simon and Schuster/New York Academy of Sciences 1988 G.
S. Paul & G. D. Leahy: Terramegathermy in the Time of the Titans:
Restoring the Metabolics of Colossal Dinosaurs, 177 – 198 in: G. D.
Rosenberg & D. L. Wolberg (ed.), R. S. Spencer ("Series Editor"): Dino
Fest. Proceedings of a conference for the general public – The Paleontological
Society Special Publication No. 7, The Department of Geological Sciences,
The University of Tennessee, Knoxville, 1994G. S. Paul: Dromaeosaurid
Archaeopteryx, www.dinosauria.com/jdp/archie/dromey.htm
(1996) G. S. Paul: Paul’s comments about Feduccia’s bird digit
paper, www.dinosauria.com/jdp/archie/paulfed.htm
(1997)J. Poling: Feathers, scutes and the
origins of birds, www.dinosauria.com/jdp/archie/scutes.htm
(1996)
J. Poling
(ed.): Responses to dinosaur lung claims, www.dinosauria.com/jdp/misc/lungs.html
(1998)
J. Poling (ed.): Skippy the dinosaur,
www.dinosauria.com/jdp/misc/scipionyx.html (1998) J. H. Reichholf: Die Vogelfeder – ein
Eiweiß-Endlager?, kosmos 4/97, 68 – 71 <Or, Why it is useful to know
German. =8-) > P. C. Sereno: A rationale for phylogenetic
definitions, with application to the higher-level taxonomy of Dinosauria,
Neues Jahrbuch für Geologie und Paläontologie Abhandlungen 210(1), 41 – 83
(October 1998)H. D. Sues: On Chirostenotes, a
Late Cretaceous oviraptorosaur (Dinosauria: Theropoda) from western North
America, JVP 17(4), 698 – 716 (December 1997) D. J.
Varricchio, F. Jackson, J. J. Borkowski & J. R. Horner: Nest and egg
clutches of the dinosaur Troodon formosus and the evolution of
avian reproductive traits, Nature 385, 247 – 250 (16 January
1997)
Xu X., Tang Z. & Wang X.: A
therizinosauroid dinosaur with integumentary structures from China, Nature
399, 305 – 354 (27 May 1999)
Xu X., Wang X. & Wu X.: A
dromaeosaurid dinosaur with a filamentous integument from the Yixian Formation
of China, Nature 401, 262 – 266 (16. 9. 1999) Zhao X.
& Xu X.: The oldest coelurosaurian [sic], Nature 394, 234f. (16 July,
1998)Despite that there are probably some
similarities, and I’d like to read it, I’ve never had the opportunity to
read
R. A. Thulborn: The avian relationships
of Archaeopteryx, and the origin of birds, because I have no
possibility to get the Zoological Journal of the Zoological Society of London
(82, 119 – 158 [1984]) anywhere near my hands <this may have changed>.
I’ve copied this reference from Feduccia (1996), and I disagree with its
cladogram reprinted there.
The extra exit for the V1 nerve in
maniraptoriforms has first been recognized in
R. T. Bakker, M. Williams & P. J. Currie:
Nanotyrannus, a new genus of pygmy tyrannosaur, from the latest
Cretaceous of Montana, Hunteria 1(5), 1 – 30, which for me is unavailable,
too <maybe no more>; a cladogram from there is reprinted in Feduccia
(1996), from where I’ve got the reference, but except that I only have a popular
article about Nanotyrannus and its V1 nerve not quoting
anything."
Oh boy, this post took me the whole day.
=8-) =8-) =8-)
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