Defining Aves: What are birds?

Dinosaur Palaeobiology and Evolution essay by Joseph Scaife 7409981

Over 9,700 species of birds fill Earth's skies today. Grouped together in the class Aves by Linnaeus (Linnaeus,
1758), this diverse and widespread group of vertebrates shares a number of features in common that separate
them from their closest living reptilian relatives: feathered wings, homeothermy, an efficient respiratory system,
an active lifestyle and more. When we look at the fossil record, however, these characteristic traits become less
diagnostic, and the line between birds and reptiles becomes blurred. So what are birds? How do we define the
class Aves?

What is a taxon?
As this is an essay about classification, it's a good idea to revise the different schools of thought in taxonomy
today. Early taxonomists have long grouped organisms by their recognisable shared traits, but often disagreed
about which traits were more fundamental to classification morphology? Reproduction? Behaviour?
Environment? Since Darwin and the advent of modern evolutionary theory, biologists have recognised that the
common descent of organisms and their placement in a universal phylogenetic tree provides a natural basis for
classification. Including evolutionary relationships in taxonomy transforms the subject from a subjective
exercise in listing and naming into a true science with predictive power.

Modern taxonomists therefore tend to fall into one of two groups, both of which use phylogeny as a basis for
classification. Phylogenetic or cladistic systematists use cladistic methods to draw up cladograms and group
organisms together that share apomorphies, i.e. derived characters inherited from a common ancestor. These
taxonomists will only accept monophyletic clades as valid biological groups, as organisms within a clade are
more closely related to each other by definition than to those outside the clade. This means that one group
cannot give rise to another distinct group (paraphyly), as the ancestral group would not form a natural clade and
would be defined by exclusion descendants are nested within parent clades, rather than separate from them.
Cladistic systematists claim their system is more natural and objective, as no external judgement is involved in
determining what defines a particular taxon once the phylogeny of that group and related groups is established.

Evolutionary or traditional taxonomists agree that phylogeny forms a natural basis for classification, but allow

paraphyletic taxa to exist, provided the descendant forms are sufficiently different from their ancestors (they
occupy a new evolutionary grade - Hickman et al. 2011, page 207) to justify their inclusion in a new taxon. To
use a simple example, take a sparrow, a crocodile, and a monitor lizard. Genetic and anatomical evidence shows
that the sparrow and crocodile share a more recent common ancestor with each other than either does with the
monitor lizard. (Fig 1a) A traditional taxonomist would note that the lizard and crocodile have a number of
features shared from their common ancestor cold blood, scaly skin and a sprawling gait, for example that
unite them together as reptiles (Reptilia). The sparrow, on the other hand, has evolved a number of characters
feathers, warm blood and so on that separate it from its reptilian relatives and justify its position in a new class
the birds (Aves). Thus, a traditional taxonomist would have no problem with the statement birds evolved from
reptiles. (Fig 1b) A phylogenetic systematist however would say the traditional taxonomist is making a
subjective value judgement in deciding which characteristics are important in classification. Objectively, the
sparrow and crocodile share a more recent common ancestor and thus belong to one clade (Archosauria); this
clade is nested within a larger clade that includes the more distantly-related monitor lizard (Diapsida). (Fig 1c)
Reptilia is invalid as a paraphyletic taxon that artificially excludes birds based on a subjective value judgement,
unless the term is expanded to include Aves and thus to become synonymous with Diapsida.

(Fig. 1 see reference in text)

These disagreements are irrelevant when it comes to defining Aves, since all definitions of Aves in common use
today refer to monophyletic clades and are thus accepted as true groups by both cladistic and tradtional
taxonomists. However, they are of importance when discussing groups such as reptiles, which are considered
paraphyletic if they exclude birds, and will be important in the final section of this essay.

Gauthier and de Queiroz (2001) in their paper on defining Aves point out that clades can be defined in two ways:
with reference to a particular common ancestor (node-and-stem based definitions, e.g. Dinosauria is the clade
descended from the last common ancestor of Passer domesticus and Triceratops horridus), or with reference to
one or more particular apomorphies (apomorphy-based definitions e.g. Dinosauria is the clade that possesses

three or more fused sacral vertebrae, a perforated acetabulum, reverse glenoid, fused ankle complex etc.) The
two types of definitions can be coincident, but do not have to be. As we shall see, this point is of particular
importance when defining the clade Avialae, a possible synonym of Aves.

Where do birds belong?

The most restrictive possible definition of birds is Neornithes, the crown group consisting of the descendants of
the last common ancestor of all living birds. The term modern birds in this essay henceforth refers to this
group.

Molecular and anatomical evidence shows that the closest living relatives of modern birds are crocodilians, and
the clade Archosauria can be defined as the group descended from the last common ancestor of these two taxa.
The presence of teeth sockets (although modern birds have lost their teeth secondarily), antorbital and
mandibular fenestrae, and a fourth trochanter on the femur are synapomorphies that can be used to distinguish
this group. Crocodilians and birds also share an efficient four-chambered heart, but since the heart does not
fossilise in extinct archosaurs and the crocodilian heart may be an adaptation for an aquatic lifestyle (Axelsson
et al.1996) it's not known whether this is an archosaurian synapomorphy too or something that has evolved
independently in birds and crocodilians.

Modern birds share a number of traits that the crocodilians do not, and that place them in more restrictive groups
within Archosauria, represented in extant fauna only by the birds but by various extinct groups:

Birds walk digitigrade with a parasaggital stance, placing them alongside the extinct pterosaurs and all
dinosaurs in the clade Ornithodira or Avemetatarsalia. This group is also called Panaves (All birds),
as it contains all animals, living or extant, closer to any given modern bird (e.g. a sparrow) than
crocodiles. Gauthier and de Queiroz (2001) list this clade as one synonym of Aves in use by zoologists,
which makes sense for extant animals, but would also bring animals such as pterosaurs, ceratopsians
and sauropods into the avian fold! Since the vast majority of zoologists and probably all
palaeontologists would be uncomfortable calling Triceratops a bird, I won't discuss this as a possible
definition of Aves any further.

Birds have a perforated acetabulum (hip socket), 11-23 fused sacral vertebrae (more than the three or

more required for defining Dinosauria), a cnemial crest and other skeletal features that mark them out
as dinosaurs, and fossil evidence shows their position within the dinosaur clade modern birds are part
of the clade Avialae, which is part of the clade Maniraptora, which is part of Coelurosauria, which is
part of Theropoda, which is part of Dinosauria. Phylogenetic systematists would insist that since birds
are descended from dinosaurs they are dinosaurs by definition, and theropods, maniraptorans and
coelurosaurs at that; and while traditional taxonomists have drawn a line between the reptilian
dinosaurs and the avian birds and placed them in two separate classes (Reptilia and Aves) (Padlan
2004 p.210-231, Hickman et al. 2011 p. 592), I shall use the term dinosaur here to refer to the clade
Dinosauria, including birds.

The question of defining Aves is therefore a question of where within the dinosaur clade the line should be
drawn between Reptilia and Aves for traditional taxonomists, or between non-avian and avian dinosaurs for
phylogenetic systematists.

Birds are endothermic

Birds have traditionally been separated from reptiles because birds are active, endothermic animals, while
reptiles are sluggish, ectothermic sprawlers. So could Aves be defined as warm-blooded archosaurs?

This essay is not the place to go into all the lines of evidence, but since the publication of Bob Bakker's The
Dinosaur Heresies in 1986 it's become increasingly accepted by palaeontologists that some, if not all, non-avian
dinosaurs were endothermic. This therefore means that Aves cannot be defined by warm-bloodedness alone,
unless we were to incorporate the rest of the dinosaurs as well. Since pterosaurs were covered in a layer of hairlike filaments known as pycnofibres, that may have been used for insulation, it's inferred that they were
endothermic too. It should be noted that crocodilians have an efficient four-chambered heart, as mentioned
earlier, associated with a high metabolism and endothermy; if this is an apomorphy of Archosauria, then perhaps
crocodiles have become secondarily ectothermic and warm blood is a characteristic of all archosaurs. No
zoologist has ever (to my knowledge) proposed that crocodilians are birds!

Other avian characteristics that crocodilians lack but other dinosaurs possess include a bird-like respiratory

system with pneumatic bones and air sacs (Sereno et al. 2008), clavicles fused to form a furcula or wishbone in
theropods (Nesbitt et al. 2009), and of course feathers.

Avifilopluma: Feathered dinosaurs

Feathers have long been considered a defining trait of birds, and are still considered a unique trait to modern
birds no other extant animals have them. Hickman et al. in their 2011 Integrated Principles of Zoology write
If an animal has feathers, it is a bird; if it lacks feathers, it is not a bird. However, we note that feathers were not
so diagnostic in the past; some theropod dinosaurs had feathers. (Hickman et al., 2011, page 591) If we retain
feathers as the single diagnostic trait of birds, and if we assume feathers only evolved once (and hence feathered
animals are a clade) then birds become synonymous with the clade Avifilopluma, the descendants of the first
archosaur with feathers homologous with those of Vultur gryphus. (Gauthier and de Queiroz, 2001)

How far back in archosaur evolution we find feathers depends on what we define as a feather. Xu and Guo
(2009) define eight stages in feather evolution, based on palaeontological evidence along with the embryonic
stages of feather development in modern birds. Foth (2011) disputes some of these stages (as found in fossils)
actually existed, and suggests some of them are crushed versions of other feather stages. Nonetheless, the
concensus among palaeontologists and developmental ornithologists is that feathers began as a single filament of
keratin, which spread out into a tuft of multiple filaments (forming a plumaceous or downy feather) growing
from a single central filament which elongated and became the central shaft or rachis in pennaceous feathers.

It's now known that pennaceous feathers existed in oviraptorosaurs, dromaeosaurs, deinonychosaurs and
avialans (about which more later), and downy feathers probably existed in all coelurosaurs (unless they were
subsequently lost, either in an entire lineage or through part of the life cycle there have been some reports of
mosaic scales in Tyrannosaurus skin impressions, for example (Gregory 2008) although whether this is actually
preserved Tyrannosaurus skin is unknown. If this claim holds up, perhaps this dinosaur lacked feathers because
feathers would cause an enormous adult Tyrannosaurus living in a warm climate to overheat although feathers
may have been possible on other parts of the body, on smaller Tyrannosaurus chicks, and filamentous feathers
are known to have existed in large adult tyrannosaurids living in cooler climates like Yutyrannus (Xu et al. 2012)

Of course, this may turn out not to be preserved Tyrannosaurus skin and perhaps the adult Tyrannosaurus was
covered in feathers). True feathers, whether through fossilised skin impressions or quill knobs on bones, have
not been identified outside Coelurosauria. Assuming they never will be found outside this group, Avifilopluma
can be defined as the clade descended from the last common ancestor of oviraptorosaurs and avialans (if we only
accept pennaceous feathers) or as synonymous with Coelurosauria (if we accept plumaceous feathers).

So are all these animals birds? Avifilopluma has been used as a definition of birds/Aves before (Gauthier and de
Queiroz, 2001; Lee and Spencer 1997) but this usage is rare nowadays as more and more feathered dinosaurs are
discovered. If Avifilopluma, Coelurosauria and Aves are synonymous, then iconic dinosaurs like Velociraptor
and Tyrannosaurus rex become classified as birds. There's no scientific reason why this would be a bad idea, but
for most people, considering dinosaurs like T. rex birds violates the idea of what a bird is, and the majority of
palaeontologists would rather drop the feathers define birds tradition than expand Aves to include all
coelurosaurs.

Moreover, there's no reason not to suspect feathers will be found in other theropod lineages, and thus
Avifilopluma may expand further. Although some non-coelosaurian theropods like Carnotarus are known to
have posessed scales (Czerkas and Czerkas, 1997), modern birds have scaly feet and so the presence of scales on
one part of the body does not preclude the possibility of feathers elsewhere. Single filaments, which Xu and Guo
(2009) define as stage 1 feathers, are known to have occurred on some ornithischian dinosaurs (Zheng et al.
2009) if these prove to be homologous with coelurosaur feathers, then the common ancestor of saurischians
and ornithischians may have posessed filamentous or plumaceous proto-feathers and Avifilopluma may end up
being synonymous with Dinosauria itself! In fact, pterosaurs are covered in a coat of hollow, probably
keratinous pycnofibres (Czerkas and Ji, 2002) if these are homologous to feathers, then perhaps Avifilopluma
would be synonymous with Avemetatarsalia. If this is true, synonymising Aves and Avifilopluma would cause
endless headaches to palaeontologists who've spent years explaining to the general public that pterosaurs are
flying reptiles and not prehistoric birds!

Avifilopluma is therefore, in my opinion, not a good clade to synonymise with Aves.

Although this essay is not the place to go into detail about hypotheses for the origin of feathers as an adaptation
(for warmth, sexual display, brooding over eggs, camouflage etc.), it's clear that feathers long predate flight in
dinosaur evolution and have been exapted by birds for flight. So if feathers don't define a bird, does flight?

Archaeopteryx and Avialae: Flying dinosaurs

Ever since its discovery in 1861, Archaeopteryx has held both the popular and scientific imagination as the first
bird - indeed, it's sometimes referred to as Urvogel, from the German first bird. The genus is now known
from 11 specimens (plus a single feather found in 1860) and two species (A. lithographica and A. siemensii)
have been identified.

Archaeopteryx does show a number of avian features, such as feathers, an opposable hallux, a fused furcula
(wishbone) and an elongate, backwards-facing pubis (superficially resembling an ornithischian dinosaur,
although birds are saurischian dinosaurs), and these characters have historically been used to classify it as a bird.
However, three of these characters have now been found in other theropods not traditionally considered birds,
and while no other dinosaurs have an opposable hallux some have a reversed hallux and there is a trend towards
the opposable hallux in the theropod lineage leading up to birds (Forster et al. 1998).

Archaeopteryx also posesses many features found in non-avian reptiles but not in modern birds, such as:

The premaxilla and maxilla are not covered by a beak, and posess teeth.

A long, bony tail that does not terminate in a pygostyle.

The ribs do not articulate with a bony sternum, as they do in modern birds, and lack uncinate proceses.
They are also thinner than typical bird ribs.

Simple concave articulation points between cervical vertebrae rather than the saddle shaped
(heterocoelus) articulation found in modern bird necks.

The pelvic girdle generally resembles that of non-avian theropods rather than birds, except for the
aforementioned reversed pubis.

There are only 6 sacral vertebrae (typical of reptiles) rather than the 11-23 seen in modern birds.

In Archaeopteryx the fibula (in the wing) and tibia (in the leg) are of roughly equal length. In modern
birds the fibula is much shorter.

In modern birds the metatarsals are fused to form the tarsometatarsus, in Archaeopteryx they remain
mostly unfused. However, in bird embryos the metatarsals are unfused, and so it's possible that
Archaeopteryx could have secondarily aborted this fusion during embryonic development.
Ceratosaurians, Avimimus and Elmisauridae all have fused tarsometatarsi, theropod groups more
distantly related to modern birds than Archaeopteryx is. So this may be a theropod apomorphy that
some groups have lost during their embryonic development, rather than an avian feature.

Archaeopteryx has gastralia, modern birds do not.

Archaeopteryx's wings terminate in a small hand made of three unfused digits with claws. No adult
bird has this, although embryos begin to develop claws, and the hoatzin retains them for much of its
juvenile stage perhaps as an adaptation for tree-climbing.

(Ostrom, 1976; de Beer (1954))

Archaeopteryx therefore appears to share more features with non-avian theropods than with modern birds.
Nonetheless, its most avian characteristic appears to have been flight. Archaeopteryx's feathers form a true, birdlike wing, with asymmetrical feathers, which could be aerofoils. The degree of asymmetry of these feathers,
when compared with the feathers of modern birds, would imply Archaeopteryx was a slow flier, rather than
flightless or a fast flier (Norberg, 1995). It has recently been discovered that the leading edge of Archaeopteryx's
feathers were melanised (Manning et al. 2013), and since melanin strengthens keratin, this could be evidence of
an adaptation to flight. Moreover, Archaeopteryx fossils are found in marine or lake sediments how did they
get there if they couldn't fly over water? Endocranial casts show that Archaeopteryx's brain is larger than most
non-avian dinosaurs but similar to that of birds, with an enlarged visual cortex, and the shape of the inner ear
preserved in the skull resembles that of modern birds more than reptiles. Archaeopteryx therefore appears to
have had a brain and sensory system adapted for flight (Witmer, 2004; Alonso et al. 2004). Archaeopteryx
lacked a bony sternum that ribs articulated to, which means flight muscles must have attached to a different bone
or to a cartilaginous sternum, making them less adapted to powered flight than modern birds are. Different
estimates of body weight, flight muscle strength and where these muscles attached have led to different authors
suggesting Archaeopteryx was either a pure glider, a glider with some limited flapping ability, or a powered flier
as modern birds are. (Senter, 2006; Nudds and Dyke 2010) In any case, the concensus among palaeontologists is
now that Archaeopteryx was a volant animal with at least some flight capability.

So where does Archaeopteryx fit in to our definition of a bird? The majority of researchers now agree that this
animal is more closely related to modern birds than to non-avian theropods (although some have classified it as a
deinonychosaur, but this has been hotly contested and is generally thought not to be supported by the evidence
(Xu et al. 2011, Senter et al. 2012) this essay will go with the majority view) and belongs to the clade Avialae
with them. This presents an interesting conundrum alluded to earlier is Avialae a node-and-stem based or
apomorphy-based clade? Some researchers define Avialae as the clade descended from the last common ancestor
of modern birds and Archaeopteryx (Padlan and Chiappe, 1997), making Archaeopteryx the most basal avialan
by definition (if we define birds as avialans, this definition has the advantage of fitting in with nostalgic views
of Archaeopteryx as the first bird!). This would be problematic if the minority view of Archaeopteryx as a
deinonychosaur turns out to be correct. Others define Avialae as those theropods closer to modern birds than
Deinonychus (Senter, 2007). Gauthier and de Queiroz (2001), however, uses an apomorphy-based definition for
Avialae the clade of flying dinosaurs. (Modern flightless birds are secondarily flightless, so they still count as
part of the flying dinosaur group, in the same way whales are still mammals although mammals are land
animals, or snakes are reptiles even though reptiles are tetrapods.) If the majority view is right about
Archaeopteryx being a flier, this would place Archaeopteryx as a basal avialan.

A few more primitive genera than Archaeopteryx are now known, such as Anchiornis, Eosinopteryx and
Aurornis (both the oldest and seemingly most basal), which appear to be closer to modern birds than
deinonychosaurs and are thus basal members of Avialae by Senter's definition (Lefvre et al. 2014) but not
Padlan and Chiappe's definition. It's not known if these could fly and thus meet Gauthier and de Queiroz's
definition of Avialae, but they possessed long feathers that covered their hind limbs and may have served as a
second pair of wings useful for flight. (Zheng et al. 2013)

It should be noted that the dromaeosaur Microraptor appears to have evolved flight independently of Avialae,
with long, asymmetrical pennaceous feathers forming an aerodynamic wing on both its arms and legs. This fourwinged dinosaur appears to have been capable of gliding and perhaps even powered flight (Dyke et al. 2013).
Dromaeosaurs are closely related to deinonychosaurs and avialans, and together they form the clade Paraves.
However, since Microraptor did not lead to modern birds, and most of its close relatives and ancestors appear to

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have been flightless (including, it's inferred, the common ancestor of Paraves), it seems more logical to call this
convergent evolution in two closely-related but distinct taxa than to expand Avialae (as flying dinosaurs) to
encompass all of Paraves because of one anomalous genus.

Trends and groups in avialan evolution

Fastovsky and Veishampel (2012) list six major steps in the evolution from the earliest Avialae to Neornithes,
and I shall list the names of clades and representative genera for each step. Direct quotes in this section are from
Fastovsky and Veishampel, page 253.

1.

A perching foot and limited flapping flight in the earliest avialans, such as Aurornis and Archaeopteryx.

2.

Reduction of the tail to form a pygostyle, a short structure at the tip of the tail formed by fusion of the
caudal vertebrae. This seems to have been a gradual process rather than a single evolutionary
development: the clade Avebrevicauda has a reduced tail, while the clade Pygostylia has a true
pygostyle. Confuciusornis, an Early Cretaceous avialan from China, has such a pygostyle, and shows
the beginning of more fused trunk vertebrae for efficient flight.

3.

Reduction in the number of trunk vertebrae, and the development of a flexible furcula, strut-like
coracoid, carpometacarpus and fully folding wings. (Fastovsky and Veishampel, page 253) These traits
define the clade Ornithothoraces (bird thorax), which includes the clade Ornithuromorpha (see stage
4) as well as the now extinct Enantiornithes (opposite birds,) a once diverse group of avialans with
fully foldable wings and gastralia

4.

Further reduction in the number of trunk vertebrae, loss of gastralia, final rotation of pubis to be
parallel with ilium and ichium. These traits, along with the development of a fully heterocoelus neck
(saddle-shaped articulation between cervical vertebrae, allowing for stability of the head; this trait
begins in basal avialans like Archaeopteryx but becomes fully modern at this point) and modern
pygostyles defines the clade Ornithuromorpha or Euronithes.

5.

[Further] reduction in the number of trunk vertebrae, decrease in the size of acetabulum [although it's
still perforated, as in all dinosaurs], formation of patellar groove... to accommodate the patella [patellae
are present in modern birds but not non-avian dinosaurs, perhaps they originated at this point?] This,
along with fully fused vertebrae (although Confuciusornis has the beginnings of a synsacrum (fused

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pelvis), this feature reaches modernity here) defines the clade Ornithurae. Notable Mesozoic
ornithurans include Ichthyornis and Hesperornis from the Late Cretaceous of the American Midwest.
6.

Loss of teeth and replacement by a keratinous beak. This is the final trait shared by all modern birds
and their common ancestor, so we've finally reached the clade Neornithes.

(Information on clades from Zhou et al. 2005, Letvre et al, 2014, and elsewhere in Fastovsky and Veishampel.)

While it is interesting to note these changes and the general trends (e.g. tail reduction, fusion of vertebrae,
rotation of the pubis, fusion of the ankle joint and an increasing forelimb length relative to hindlimb length) that
accompanied them, no authors I've found have used any of them to define birds/Aves. The term birds or
Aves usually refers to either the whole clade Avialae (flying dinosaurs) or Neornithes (toothless, beaked
modern birds) alone, and never anything inbetween.

It is interesting to note that Neornithes were the only group of dinosaurs to survive the K-Pg mass extinction,
and therefore the statement all non-avian dinosaurs died out at the K-Pg extinction is true whether you define
Aves as Avialae or Neornithes (with the possible exception of Qinornis paleocenica, a Palaeocene avialan that
may or may not have been non-neoornithine and thus, if Aves is restricted to Neornithes, may be the only nonavian dinosaur known to have survived the K-Pg extinction (Xue, 1995; Longrich et al, 2011)).

So what are birds?

In this essay we have noted along with Gauthier and de Queiroz (2001) four possible definitions of Aves, along
with alternative clade names for each. From most to least exclusive:

Aves could refer to all archosaurs closer to modern birds than to crocodiles, a clade also called Panaves,
Avemetatarsalia or Ornithodira and which would include all dinosaurs and pterosaurs as part of the
avian group.

Aves could refer to all feathered dinosaurs, a clade also called Avifilopluma and which may be
synonymous with Coelurosauria (or even larger groups). Although this would keep the traditional
feathers are diagnostic of birds criterion, both of the first two definitions would include many
dinosaurs traditionally described as non-avian, such as Tyrannosaurus, and are thus not popular among
palaeontologists or the general public.

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Aves could refer to all flying feathered dinosaurs (or all closer to modern birds than to
deinonychosaurs, or all those descended from the last common ancestor of modern birds and
Archaeopteryx.) This clade is also known as Avialae.

Aves could refer to everything descended from the last common ancestor of all living birds, defined by
their toothless beaks and also known as the clade Neornithes.

Although Gauthier and de Queiroz list a few more definitions they have found in the literature (e.g. Aves =
Theropoda (Thulborn, 1975)), these four are the most common, and only the last two make sense. So should
Aves be synonymous with Avialae or Neornithes?

This is where the disagreements between phylogenetic systematics and evolutionary taxonomy come into play. A
phylogenetic systematist would only accept monophyletic clades, and thus, both Avialae and Neornithes are
valid taxa. However, this form of taxonomy is incompatible with the traditional Linnean ranks, as one group
cannot give rise to a separate group of equal rank (as opposed to a lesser group nested within it). Since the term
Aves was invented by Linnaeus as a class in 1758, there seems no good reason to retain it within a cladistic
classification system anymore. Avialae and Neornithes are less ambiguous terms, and neither constitutes a class
in a system where taxonomic ranks are obsolete! Aves would take priority over either of these names, as an older
taxon name, if it could be shown to be synonymous with one of them but taxonomists cannot agree which
clade Aves is synonymous with. Therefore I think the best solution in a cladistic system would be to drop Aves
as a formal taxon, retain the word bird only as an ambiguous, informal term, and keep Avialae and Neornithes
as clades.

In traditional taxonomy, phylogeny is combined with the traditional Linnean ranks and paraphyly is acceptable
when a descendant taxon is judged to be different enough from its ancestors to constitute a new evolutionary
grade (Hickman et al. 2011, page 207). Birds - agile, feathered, endothermic fliers - certainly seem to justify
separation from their ancestral taxon, the sluggish, ectothermic, scaly and sprawling Reptilia, and so I'm in
favour of retaining the traditional taxon name of Aves for a class in such a system. But where do we draw the
line? Most taxonomists of both schools now use Aves as a synonym for Neornithes (Gauthier and de Queiroz
2001; Hickman et al. 2011, chapter 27), although some use Avialae (Godefroit et al. 2013, Zhou 2005 also

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defines Enantiornithes as part of Aves). In my opinion, though, the characteristic that defines Neornithes
toothlessness is relatively minor and does not justify its separation as a new class. Avialae, as the group of
flying, feathered dinosaurs, seems to me to be a more logical clade for traditional taxonomists to synonymise
with Aves, as these animals are adapted to flight in the air rather than locomotion on land, and this new mode of
life seems to me to be a far more major and fundamental change than posessing a toothless beak a change
worthy of a new class, analogous to the way fish became tetrapods when they left the water. An alternative is to
remove Aves as a class and include Avialae (and thus Neornithes) within the class Reptilia, which would sit well
with cladists as it makes Reptilia a monophyletic group (synonymous with Diapsida, or with Sauropsida if
anapsids are counted. Reptilia would still be paraphyletic if the synapsid reptiles such as pelycosaurs were
included, though, because synapsid reptiles gave rise to the class Mammalia.) Most zoologists, however, aren't
prepared to call birds reptiles cladists use the term as an informal group while traditional taxonomists accept it
as a valid paraphyletic class, both uses excluding birds. (Tudge, 2000, chapter 21).

Bob Bakker in the final chapter of The Dinosaur Heresies (1986) proposes a much more radical solution to this
problem. If we accept reptiles are paraphyletic and birds evolved from them, we can see that the major traits that
differentiate modern birds from modern reptiles homeothermy, feathers, an upright stance and an active
lifestyle, whether on the ground or in the air emerged far further back in archosaurian evolution than either
Avialae or Neornithes, and thus birds aren't the only animals that Bakker thinks should be excluded from a
paraphyletic Reptilia. The class Aves is dead long live the class Dinosauria!

When all is said and done, however, these arguments are purely academic. Correct nomenclature is important,
and when scientists around the world use the term Aves other scientists should know what they mean by that
term. But Avialae and Neornithes already exist as unambiguous clade names, and deciding which of the two (or
any other relevant group) gets to be called Aves is a matter more of opinion than science. As Richard Feynman
said, You can know the name of that bird in all the languages of the world, but when you're finished, you'll
know absolutely nothing whatsoever about the bird. You'll only know about humans in different places, and what
they call the bird. ...I learned very early the difference between knowing the name of something and knowing
something. All neoornithines.. .all avialans... and all dinosaurs are fascinating creatures, no matter what we
wish to call them. What are birds? We just don't know.

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