Microraptor: not a ‘raptor’??

Earlier
the large reptile tree nested two putative dromaeosaurs with composgnathid/tyrannosaurs, Tianyuraptor and Zhenyuanlong. Today the famous four-wiinged dinosaur/bird Microraptor (Figs. 1, 2) is added to that list, nesting between Compsognathus and Tianyuraptor, all three basal to T-rex.

Figure 1. Microraptor gui (IVPP V 13352) shown in two photos and with DGS tracing of bones and feathers.

Figure 1. Microraptor gui (IVPP V 13352) shown in two photos and with DGS tracing of bones and feathers. Click to enlarge.

This is the specimen
that inspired a PBS Nova special and a race between competing teams of paleontologists to figure out the best usage for the odd foot feathers. The Kansas team led by Dr. Larry Martin produced a sprawling model that went against everything we know di dinosaur hind limbs.

Figure 2. Microraptor gui (IVPP V 13352) reconstructed from tracings in figure 1. There are no surprises here, except a provisional closer relationship with Compsognathus than with Velociraptor. Microraptor has a large pedal claw two, but it is not quite the killing claw seen in droamaeosaurs.

Figure 2. Microraptor gui (IVPP V 13352) reconstructed from tracings in figure 1. There are no surprises here, except a provisional closer relationship with Compsognathus than with Velociraptor. Microraptor has a large pedal claw two, but it is not quite the killing claw seen in droamaeosaurs.

So this makes three former dromaeosaurs
now nesting with long-legged Compsognathus and Tyrannosaurus. Among them, only Microraptor has long arms/wings. Zhenyuanlong has equally substantial feathers. So this adds credulity to the idea that Compsognathus was well feathered. Only Microraptor has a posteriorly directed pubic foot, but see Compsognathus (Fig. 4) for its derivation. This is not a posteriorly directed pubis.

Figure 4. Compsognathus was not preserved with feathers, but with a sister taxon like Microraptor, it might have had substantial feathers.

Figure 4. Compsognathus was not preserved with feathers, but with a sister taxon like Microraptor, it might have had substantial feathers.

Is Microraptor a bird (clade Aves)?
Wikipedia (Evolution of Birds) defined Aves as “all descendants of the most recent common ancestor of a specific modern bird species (such as the house sparrow, Passer domesticus), and either Archaeopteryx, or some prehistoric species closer to Neornithes (to avoid the problems caused by the unclear relationships of Archaeopteryx to other theropods).[ If the latter classification is used then the larger group is termed Avialae. Currently, the relationship between dinosaurs, Archaeopteryx, and modern birds is still under debate.”

Is Microraptor a member of the clade Avialae?
Wikipedia defines the clade Avialae “a clade of dinosaurs containing their only living representatives, the birds. It is usually defined as all theropod dinosaurs more closely related to modern birds (Aves) than to deinonychosaurs, though alternate definitions are occasionally used (see below).” 

So, Microraptor is not a bird. 
In the same light, not all Archaeopteryx specimens are birds, but Wellnhoferia (aka The Solnhofen specimen, Archaeopteryx grandis) apparently is a bird as it nests closest to living birds of all Solnhofen specimens.

Yes
I don’t have a complete list of theropods in the large reptile tree. But this is what the tree recovers at present. If valid, theropods with long feathers on their forelimbs appear earlier than some workers think. And maybe I’m just catching up to the rest of them.

There are other specimens out there referred to Microraptor
and I have not tested them yet. Perhaps one or more are more closely related to Velociraptor. 

Addendum
Here is the skull of the QM V 1002 specimen of Microraptor (Fig. 5, Xing et al. 2013). The two nest together in the large reptile tree, but differ in several traits. They are not conspecific.

Figure 5. The skull of another Microraptor, QM V1002. The two nest together in the large reptile tree.

Figure 5. The skull of another Microraptor, QM V1002, the fish eater. The two nest together in the large reptile tree. I’m a little confused by the occiput. I’ll get back to that later.

References
Xing L, Persons WS, Bell PR, Xu X, Zhang J-P, Miyashita T, Wang F-P and Currie P 2013. Piscivery iin the feathered dinosaur Microraptor. Evolution 67(8):2441-2445.
Xu X, Zhou Z, Wang X, Kuang X, Zhang F, and Du X 2003. Four-winged dinosaurs from China. Nature, 421: 335–340.

wiki/Microraptor

 

 

Intermedium reappears in birds

Ossa-Fuentes L, Modozis J and Vargas AO 2015
discover a detail of interest in bird osteology and ontogeny.

They report, “This work has revealed that the ascending process [of the ankle] does not develop from either the heel bone or the ankle bone, but from a third element, the intermedium. In the ancient lineage of paleognath birds (such as tinamous, ostriches and kiwis) the intermedium comes closer to the anklebone, producing a dinosaur-like pattern. However, in the other major avian branch (neognaths), which includes most species of living birds, it comes closer to the heel bone; that creates the impression it is a different structure, when it is actually the same.”

And that’s not all… They continue: “More remarkably, however, this finding reveals an unexpected evolutionary transformation in birds. In embryos of the land egg-laying animals, the amniotes (which include crocodilians, lizards, turtles, and mammals, who secondarily evolved live birth) the intermedium fuses to the anklebone shortly after it forms, disappearing as a separate element. This does not occur in the bird ankle, which develops more like their very distant relatives that still lay their eggs in water, the amphibians. Since birds clearly belong within landegg-laying animals, their ankles have somehow resurrected a long-lost developmental pathway, still retained in the amphibians of today – a surprising case of evolutionary reversal. The study also presented fossil evidence from juvenile specimens of toothed birds from the Cretaceous period. These show that, at this early stage of bird evolution, the ascending process already developed separately.”

On a similar note,
as you may recall from this earlier blog post, the pre-amniote, and almost pre-tetrapod, digit zero, the manual digit medial to the thumb, which is absent in almost all derived tetrapods, also appears on Limusaurus and caused the phase shift confusion noted earlier.

References
Ossa-Fuentes L, Modozis J and Vargas AO 2015. Bird embryos uncover homology and evolution of the dinosaur ankle. Nature Communications. DOI: 10.1038/natcomms9902
Diaz RE & Trainor PA 2015. Hand/foot splitting and the ‘re-evolution’of mesopodial skeletal elements during the evolution and radiation of chameleons. BMC evolutionary biology, 15(1), 184.
https://paleobiologia.wordpress.com (blog en español)
http://www.nature.com/ncomms/2015/151113/ncomms9902/full/ncomms9902.html

Walking around like a chicken with its head cut off

Since chickens are dinosaurs,
one wonders if such stories (see below) ever replayed in the Mesozoic after a predator encounter.

Figure 1. Mike the headless chicken, circa 1945. Click for webpage.

Figure 1. Mike the headless chicken, circa 1945. Click for webpage.

I happened to be listening to a story on NPR (National Public Radio) today
about a chicken surviving three days with its head cut off, and memories of my time as a kid on a Nebraska farm seeing the same, inspired today’s blog.

Here’a a YouTube video of a headless chicken for the morbidly curious, apparently from China.

And here’s another video about Mike, the headless chicken, back in the 1940s. He lived for, they say, 18 months after his decapitation. And here is another video about Mike. There’s a Wikipedia article here. And there’s even a website devoted to Mike where you can get T-shirts and other such memorabilia if you you are so inclined.

Apparently if you cut too high,
part of the brainstem and an inner ear can remain connected to the nervous system. So the chicken can maintain balance, react to sounds and have no head.

 

 

 

Rhea finger claws + Star Wars flashback

I’m digging into my illustration vaults today…
as I try to dash out a blog while working on a more involved and serious piece that will appear here shortly.

Several decades ago
I had the all the excitement every novice feels while getting into a new line of interest, in this case, paleontology. To that end, I had a frozen rhea (Rhea americana) sent from a farm. I processed and mounted the skeleton. Along the way I was surprised to find the surgically sharp finger claw on manual digit 1. This comes as no surprise to ornithologists or paleontologists, but the image of the Rhea skeleton was used in a 1996 book I illustrated for author, Don Lessem, “Raptors – The Nastiest Dinosaurs.”

Figure 1. Partial skeleton of Velociraptor compared to similar bones in the living bird, Rhea americana, photographed from a skeleton I processed several decades ago. This illustration was originally published in Don Lessem's "Raptors, the Nastiest Dinosaurs."

Figure 1. Partial skeleton of Velociraptor compared to similar bones in the living bird, Rhea americana, photographed from a skeleton I processed several decades ago. This illustration was originally published in Don Lessem’s “Raptors, the Nastiest Dinosaurs.”

 

On a lighter note,
with the latest Star Wars movie (The Force Awakens) about to appear (Dec. 18), I’m dragging out an old illustration from 1982, when Time Magazine reported the third movie in the original trilogy from George Lucas would be called, “Revenge of the Jedi” (Fig. 2). As everyone knows, when the movie was released the title became “Return of the Jedi” (Fig. 3).

At one time the third Star Wars movie was going to be called, Revenge of the Jedi. This was one of my first illustrations, was never submitted, but was used as my calling card back in the day (circa 1982-1986), prior to GIANTS and the genesis of my interest in paleontology.

Figure 2. At one time the third Star Wars movie was going to be called, Revenge of the Jedi. This was one of my first illustrations, was never submitted, but was used as my calling card back in the day (circa 1982-1986), prior to GIANTS and the genesis of my interest in paleontology. Fuzziness comes from fingerprint on my phone lens.

Official poster for Return of the Jedi. If you look closely Darth Vader will appear in the deep indigo sky.

Figure 2. Official poster for Return of the Jedi. If you look closely Darth Vader will appear in the deep indigo sky.

As a novice illustrator
I was inspired to create a poster for ROTJ which I ultimately used on my business card to advertise my availability (while shamelessly glomming on to something larger and more magnificent). The type font is original. Luke is lit on his right by red light, representing the dark side, under the strong influence of Darth Vader. The light saber lights his left as he, in my naiveté, powerfully grabs the light saber.

Only several years later
did the idea for the book GIANTS cross my mind. The rest is pre-history.

Thank you
for putting up with this little indulgence.

 

Liaoning bird embryo IS a Chinese Archaeopteryx

Updated 11/22/2015 with high rez data sent by Dr. Zhou. A new analysis nests the embryo with the holotype Archaeopteryx lithographica, the London specimen, a basal enantiornithine bird. 

Zhou and Zhang (2004)
described a small, precocial, final stage bird embryo from the Liaoning Province (Early Cretaceous, 121mya, IVPP V14238). Strangely, no eggshell was preserved (Fig. 1), but the tucked shape of the embryo indicated that it had not yet hatched. Northern China was a forested landscape dominated by active volcanoes and sprinkled with lakes and streams at the time. No adults were closely associated, but enantiornithine birds are common in that formation.

Figure 1. Click to enlarge. Liaoning bird embryo IVPP V14238 reconstructed Egg tracing in DGS compared to original tracing (in olive). Note the universally observed long tail and the continuation of the tail vertebrae past the back of the skull. Note the broken clavicles. When rotated they form more of a U shape. The dorsal coracoid is a convex and the ventral scapula is concave, an enanthiornithine key trait.

Figure 1. Click to enlarge. Liaoning bird embryo IVPP V14238 reconstructed Egg tracing in DGS compared to original tracing (in olive). Note the universally observed long tail and the continuation of the tail vertebrae past the back of the skull. Note the broken clavicles. When rotated they form more of a U shape with appropriate spacing of the coracoids. The dorsal coracoid is a convex and the ventral scapula is concave, an enanthiornithine key trait.

The Zhou and Zhang Abstract
“An embryo of an enantiornithine bird has been recovered from the Lower Cretaceous rocks of Liaoning, in northeast China. The bird has a nearly complete articulated skeleton with feather sheet impressions and is enclosed in egg-shaped confines. The tucking posture of the skeleton suggests that the embryo had attained the final stage of development. The presence of well-developed wing and tail feather sheets indicates a precocial developmental mode, supporting the hypothesis that precocial birds appeared before altricial birds.”

Figure 2. The Liaoning bird egg IVPP V14238 in situ with DGS tracing in color. This hirez version updates a prior lo rez version. Length of shell is 3.5 cm.

Figure 2. The Liaoning bird egg IVPP V14238 in situ with DGS tracing in color. This hirez version updates a prior lo rez version. Length of shell is 3.5 cm.

Zhou and Zhang 
did not create a reconstruction (Fig.1) nor attempt to untuck the embryo. Bird embryos shift into a tuck position before hatching as they begin to occupy most of the egg. No egg tooth is present on this specimen.

Figure 3. The Liaoning embryo compared to its closest sister, the London specimen of Archaeopteryx (holotype). The egg is the correct size to pass through the ischia if they were separated distally. like modern birds,

Figure 3. The Liaoning embryo compared to its closest sister, the London specimen of Archaeopteryx (holotype). The egg is the correct size to pass through the ischia if they were separated distally. like modern birds,

Zhou and Zhang report [with my observations in brackets]:
“The embryo has several enantiornithine apomorphies such as a strutlike coracoid with a convex lateral margin [yes], a V-shaped furcula [maybe], metacarpal III extending well past metacarpal II distally  [no], and metatarsal IV being more slender than metatarsals II or III [no]. My observations were improved with a high resolution image (Fig. 2). The Liaoning embryo nests with the holotype Archaeopteryx (London specimen), which nests at the base of the Enantiornithes.

This is the first
Cretaceous avian embryo preserved with feathers, sheathed, not open vanes. These indicate the embryo was precocial, able to move and feed independently shortly after hatching. This specimen demonstrates that the genus Archaeopteryx survived into the Early Cretaceous.

Figure 4. The Liaoning embryo bird nests with several Archaeopteryx specimens in the large reptile tree, AND with enanthiornithes. The large reptile tree does not specifically test for the classic enantiornithine traits, but correctly nested the embryo with adult enantiornithines.

Figure 4. The Liaoning embryo bird nests with several Archaeopteryx specimens in the large reptile tree, AND with enanthiornithes. The large reptile tree does not specifically test for the classic enantiornithine traits, but correctly nested the embryo with adult enantiornithines.

Compare this bird embryo to a precocial pterosaur embryo or three
like Pterodaustro, the IVPP embryo or the JZMP embryo. Embryo pterosaurs have the proportions of an adult. They grow isometrically. Hatchling birds, like the Liaoning embryo, had juvenile proportions with a large head, short tibia and short metatarsus. They grew allometrically, but not as allometric as living altricial (helpless) bird hatchlings.

“Several previously known theropod embryos and the late Cretaceous avian embryos all seem to be preocial animals, judged purely from skeletal evidence,” Zhou said.

Nat Geo
reported, “Zhou said several other enantiornithine species are known from the deposit where the latest fossil was found, but that it was difficult to link the embryo to a specific genus or species.” Unfortunately Zhou and Zhang eyeballed the embyro. They did not attempt a phylogenetic analysis (Fig. 4).

Kevin Padian
quoted in NatGeoOnline noted that half of the fossil’s characteristics are not exclusive to enantiornithines. He added that characteristics that would identify the fossil an enantiornithine are “either dubious or not well preserved on the specimen. But then, what else could it be?” Padian asked. I agree, but then neither of us has seen the fossil first hand.

Figure 4. Enanthiornithine birds to scale. Click to enlarge.

Figure 4.  A selection of Enanthiornithine birds to scale. None of these nest closer to the Liaoning embryo. These taxa all have a shorter tail and a more gracile clavicle and other traits listed in the large reptile tree.

Others have warned me
that juveniles and embryo reptiles, like pterosaurs and tritosaurs, cannot be added to phylogenetic analyses because they tend to nest with other adults*. Actually I’d like to see that happen. At present I’m a skeptic. This was a test of that hypothesis, but it was done with a precocial embryo with a relatively larger head, shorter neck and shorter limbs. I don’t see the problem with adding this embryo (Fig. 1) or precocial pterosaur embryos to analyses. But I’m willing to listen to good arguments with valid data.

*Bennett (2006) considered small adult pterosaurs as juveniles of larger germanodactylids based on long bone lengths rather than phylogenetic analysis. Eyeballing, charts and clouds of data points are no replacements for reconstructions and phylogenetic analysis. Hope you agree…

If this is an enantiornithine
which one is it most like? Archaeopteryx lithographica.

If this is an archaeopterygid
we now have some more ontogenetic clues and patterns to work with. You can see (Fig. 1) which body parts get larger and which get smaller during maturation.

Actually it’s both!

References
Bennett SC 2006. Juvenile specimens of the pterosaur Germanodactylus cristatus, with a review of the genus. Journal of Vertebrate Paleontology 26:872–878.
Zhou Z and Zhang F-C 2004. A Precocial Avian Embryo from the Lower Cretaceous of China. BREVIA Science 22 October 2004: 306 no. 5696 p. 653. DOI: 10.1126/science.1100000. online abstract here

NatGeoOnline

What makes a bird a bird? Everyone knows, it’s not feathers any more…)

The line between birds and theropod dinosaurs
has become increasingly fuzzy now that so many non-birds have feathers and other former bird-only traits.

This is a good sign
that evolutionary theory embraces: small changes and a gradual accumulation of traits in derived taxa.

Ultimately
it may come down to a single defining trait (like mammary glands in mammals, or alternatively a squamosal/dentary jaw joint when soft tissue is missing) when you have lots of taxa near the base of a new major clade. So what is that trait? Or what are those traits as recovered by the large reptile tree?

The basal bird and its proximal outgroup
At present the last common ancestor of all extant birds, scansoriopterygids and enantiornithes in the large reptile tree. is the Thermopolis specimen of Archaeopteryx (Fig. 1). The original authors (Mayr et al. 2007; Rauhut 2013) did not employ a phylogenetic analysis, so perhaps did not realize what they had.

For now
the pre-bird theropod, Eosinopteryx (Fig.1) nests just basal to the basal bird theropod, Archaeopteryx. You might find it interesting to see which traits differentiate the latter from the former in the large reptile tree. This list, short as it is, is by no means complete. It simply reflects the general characters used for all reptiles in the large reptile tree.

Figure 1. Eosinopteryx, a pre-bird, compared to Archaeopteryx, a basal bird to scale. Click to enlarge.

Figure 1. Eosinopteryx, a pre-bird, compared to Archaeopteryx, a basal bird to scale. Click to enlarge.

Archaeopteryx (Thermopolis) novelties vs. Eosinopteryx

  1. Frontal/parietal suture straight and > than frontal/nasal suture
  2. Metacarpals 2-3 subequal
  3. Pubis and ischium oriented posteriorly (convergent with some deinonychosaurs)
  4. Pedal 4 subequal to metatarsal 4  (convergent with some deinonychosaurs)
  5. Pedal 2.1 not > p2.2
  6. Metatarsal 5 shorter than pedal digit 5 (all vestigial, of course)
Figure 2. The coracoid of the Thermopolis specimen is not as elongate as in the more derived taxa. It is just barely not a disc. Thus, this basal taxon was not quite the flapper as the other Solnhofen birds.

Figure 2. The coracoid of the Thermopolis specimen is not as elongate as in the more derived taxa. It is just barely not a disc. Thus, this basal taxon was not quite the flapper as the other Solnhofen birds.

Unfortunately
none of these traits are unique to the bird clade.

I thought, perhaps
that an elongate and locked down coracoid (the key to the origin of flapping) would prove to be present in all basal birds. Such a coracoid is indeed present in other specimens of Solnhofen birds, but not in the Thermopolis specimen (Fig. 2), the basalmost example. 

So what we are seeing
in these six Solnhofen birds are discrete steps in the evolution of the flapping behavior, necessary for creating thrust and ultimately flight, as in many living birds. Just as in Late Jurassic pterosaurs, the island/lagoon environment of Solnhofen was as powerful an agent as the Galapagos islands at splitting basal birds into various clades.

From the Mayr et al. abstract on the Thermopolis specimen:
“We describe the tenth skeletal specimen of the Upper Jurassic Archaeopterygidae. The almost complete and well-preserved skeleton is assigned to  Archaeopteryx siemensii
 Dames, 1897 and provides significant new information on the osteology of the Archaeopterygidae. As is evident from the new specimen, the palatine of Archaeopteryx
 was tetra-radiate as in non-avian theropods, and not triradiate as in other avians. Also with respect to the position of the ectopterygoid, the data obtained from the new specimen lead to a revision of a previous reconstruction of the palate of Archaeopteryx. The morphology of the coracoid and that of the proximal tarsals is, for the first time, clearly visible in the new specimen. The new specimen demonstrates the presence of a hyperextendible second toe in Archaeopteryx*.  This feature is otherwise known only from the basal avian Rahonavis and deinonychosaurs (Dromaeosauridae and Troodontidae), and its presence in Archaeopteryx provides additional evidence for a close relationship between deinonychosaurs and avians**. The new specimen also shows that the first toe of Archaeopteryx was not fully reversed but spread medially, supporting previous  assumptions that Archaeopteryx was only facultatively arboreal*. Finally,we comment on the taxonomic composition of the Archaeopterygidae and conclude that Archaeopteryx bavarica Wellnhofer, 1993 is likely to be a junior synonym of  A. siemensii****, and Wellnhoferia grandis Elzanowski, 2001 a junior synonym of  A. lithographica***** von Meyer, 1861.”

* Actually not as prominent as in deinonychosaurs. Such a toe works just as well at climbing tree trunks as climbing dinosaur flanks.

**This may be a convergence as the two clades are separated by taxa without a hyper extensible pedal 2.

*** Perhaps facultatively able to perch, but arboreality would have been a precursor behavior.

**** These two are sisters in the large reptile tree.

***** These two are not sisters.

Other traits in the Theromopolis specimen 
visible in Figure 1 not present in the large reptile tree include the following:

  1. Smaller antorbital fenestra
  2. Longer attenuate tail
  3. Slightly narrower coracoids
  4. Slightly larger forelimb
  5. Bowed gap between ulna and radius
  6. More gracile pubis, posteriorly oriented
Figure 3. Archaeopteryx Thermopolis pedal digit 2 (in pink). Pedal 2.2 was capable of hyperextension (see figure 4).

Figure 3. Archaeopteryx Thermopolis pedal digit 2 (in pink). Pedal 2.2 was capable of hyperextension (see figure 4).

Mayr et al. looked at pedal digit 2
and noticed it was capable of hyperextension (Fig. 3). They likened it to pedal digit 2 in deinonychosaurs (Fig. 4) which is famous for its ability to elevate the ‘killer claw’.

Figure 4. Deinonychus with elevated pedal digit 2 demonstrating hyperextension.

Figure 4. Deinonychus with elevated pedal digit 2 demonstrating hyperextension.

The large reptile tree
does not nest birds with deinonychosaurs. Rather Xiaotingia and Eosinopteryx nest between these clades. And Xiaotingia also has a similar pedal 2.1 (Fig. 5).

Figure 5. Pedal digit 2 in Xiaotiniga shows the ability to hyperextend pedal 2.2.

Figure 5. Pedal digit 2 in Xiaotiniga also shows the ability to hyperextend pedal 2.2.

On a final note:
Mayr et al. (2007) report four premaxillary teeth in the Thermopolis specimen. I think they might have missed counting the anteriormost premaxillary tooth (Fig. 6) bringing the total to five.

Figure 6. Archaeopteryx, Thermopolis specimen, premaxilla with five teeth, not four, identified here.

Figure 6. Archaeopteryx, Thermopolis specimen, premaxilla with five teeth, not four, identified here.

References
Rauhut OWM 2013. New observations on the skull of Archaeopteryx. Paläontologische Zeitschrift 88(2)211-221.
Mayr G, Pohl, B, Hartmann S and Peters DS 2007. The tenth skeletal specimen of Archaeopteryx. Zoological Journal of the Linnean Society 149:97-116.

Archaeopteryx: the scattered skull of the London specimen

Updated November 8, 2015 with cranial data from Alonso 2004 that I just became aware of.

Figure 1. Archaeopteryx London specimen skull. The anterior is more clearly presented and has been previously illustrated. Here I colorized matrix discontinuities that could be posterior skull elements. At least they all fit together in a basic Archaeopteryx-type skull that matches other specimens.

Figure 1. GIF animation of the skull of the London specimen of Archaeopteryx. Perhaps other bones are also present. If so I did not identify them. The bones here are clear, less clear and not very clear. Compare these colors to the colors in the reconstruction and you’ll see a close correspondence to the bones of other specimens.

As far as I know,
prior workers did not identify or illustrate the posterior skull bones of the London specimen of Archaeopteryx (Fig. 1, but see below). Bones left only the faintest of impressions (if correct here), but seem to correspond to the same bones of better known specimens (Fig. 4). Higher resolution images should confirm or refute these tracings.

New data (November 8, 2015)
came in the form of Alonso et al. 2004, which extricated and CT scanned the skull of the London Archaeopteryx. The new illustration in figure 2 reflects that data. Apologies that I was not aware of this at the time of this first posting.

Figure 2. A new paper (Alsonso et al. 2004) on the cranium of this specimen has come to my attention. The cranium was buried in the matrix and these new illustrations reflect the more complete data.

Figure 2. A new paper (Alsonso et al. 2004) on the cranium of this specimen has come to my attention. The cranium was buried in the matrix and these new illustrations reflect the more complete data.

Every bone here
appears to fit and not stray too far from morphologies established by better preserved skulls. As noted earlier, the large number of premaxillary teeth in the London specimen, along with other traits, make it distinct from the Eichstaett specimen (Figs. 3, 4).

While we’re on the subject of basal birds,
here are a few to scale (Figs. 3, 4). It is notable that the more primitive ones are the smaller ones in this selection of taxa.

Figure 4. Enanthiornithine birds to scale. Click to enlarge.

Figure 4. Enanthiornithine birds to scale. Click to enlarge. Evidently there are a few other taxa without a sternum in this clade.

Be sure to click on figure 4 to see it at full size.
The stem birds, Xiaotingia and Eosinopteryx form a short-face clade with their own autapomorphies. Rahonavis nests with Velociraptor, not with birds in the large reptile tree.

Figure 4. Archaeopteryx and a few stem birds to scale compared to a chicken (Gallus). Click to enlarge.

Figure 4. Archaeopteryx and a few stem birds to scale compared to a chicken (Gallus). Click to enlarge.

The convergence of Late Jurassic birds and Late Jurassic pterosaurs
Here it is clear that the reduction of the long tail in birds occurred with phylogenetic miniaturization and neotony. Earlier I demonstrated the same tail reduction in four clades of pterosaurs that ultimately developed ‘pterodactyloid’-grade traits. They each had their genesis in tiny pterosaurs experiencing phylogenetic miniaturization and neotony.

The refusal of pterosaur workers
to recognize that embryo and juvenile pterosaurs match their parents, and that tiny Solnhofen pterosaurs are adults the size of living hummingbirds is the reason why their cladograms fail to demonstrate gradual accumulations of traits in derived taxa. Odd that tiny birds get novel generic names, but tiny pterosaurs do not.

It may be
that only tiny birds survived the end of the Jurassic, just like tiny pterosaurs. Later they both developed into larger forms.

Rahonavis
(Forster et al. 1998) survived into the Latest Cretaceous (Maastrichtian). Not sure whether it stayed small or evolved smaller than other velociraptors. At present it nests basal to that clade.

I still think reconstructions bring necessary data to the table. 
Hope you do too.

References
Alonso PD, Milner AC, Ketcham RA Cookson MJ and Rowe TB 2004. The avian nature of the brain and inner ear of Archaeopteryx. Nature 430:666-669.
Forster CA, Scott D, Chiappe LM, Krause DW. 1998. The Theropod Ancestry of Birds: New Evidence from the Late Cretaceous of Madagascar. Science 279 (5358): 1915–1919.