Current Biology 22, 2262–2267, December 4, 2012 ª2012 Elsevier Ltd All rights reserved
052
Primitive Wing Feather Arrangement
in Archaeopteryx lithographica
and Anchiornis huxleyi
Nicholas R. Longrich,1,* Jakob Vinther,2,3 Qingjin Meng,4
Quangguo Li,4 and Anthony P. Russell5
1Department of Geology and Geophysics, Yale University,
PO Box 208109, New Haven, CT 06520-8109, USA
2Jackson School of Geosciences, University of Texas at
Austin, 1 University Station C1110, Austin, TX 78712, USA
3Departments of Earth and Biological Sciences, University
of Bristol, Woodland Road, Bristol BS8 1UG, UK
4Beijing Museum of Natural History, 126 Tianqiao South Street,
Beijing 100050, China
5Department of Biological Sciences, University of Calgary,
2500 University Drive NW, Calgary, AB T2N 1N4, Canada
Summary
In modern birds (Neornithes), the wing is composed of
a layer of long, asymmetrical flight feathers overlain by short
covert feathers [1–3]. It has generally been assumed that
wing feathers in the Jurassic bird Archaeopteryx [4–9] and
Cretaceous feathered dinosaurs [10, 11] had the same
arrangement. Here, we redescribe the wings of the archaic
bird Archaeopteryx lithographica [3–5] and the dinosaur
Anchiornis huxleyi [12, 13] and show that their wings differ
from those of Neornithes in being composed of multiple
layers of feathers. In Archaeopteryx, primaries are overlap-
ped by long dorsal and ventral coverts. Anchiornis has
a similar configuration but is more primitive in having short,
slender, symmetrical remiges. Archaeopteryx and Anchior-
nis therefore appear to represent early experiments in the
evolution of the wing. This primitive configuration has
important functional implications: although the slender
feather shafts of Archaeopteryx [14] and Anchiornis [12]
make individual feathers weak, layering of the wing feathers
may have produced a strong airfoil. Furthermore, the layered
arrangement may have prevented the feathers from forming
a slotted tip or separating to reduce drag on the upstroke.
The wings of early birds therefore may have lacked the range
of functions seen in Neornithes, limiting their flight ability.
Results
The wing feathers of Archaeopteryx lithographica are best
seen in the counterslab of the Berlin specimen, HMN (Hum-
boldt Museum für Naturkunde) 1880 (Figures 1 and 2). Previ-
ously, it was shown that the feathers are not preserved as
simple impressions lying in a single plane but instead repre-
sent collapsed molds passing through the matrix three-dimen-
sionally [4] (see Figure S1 available online). This insight is
critical to understanding Archaeopteryx, because where the
split in the matrix passes through different layers of sediment,
it is possible to examine feathers lying in different planes [4].
Although the split slab primarily exposes the ventral surface
of the wing, in places the ventral feathers are broken away to
expose feathers lying more dorsally.
*Correspondence: nicholas.longrich@yale.edu
http://dx.doi.org/10.1016/j.cub.2012.09.
Report
Along the margin of the wing, eight primaries are visible. The
wing’s leading edge is formed by four successively longer
primaries (P5–P8); their vanes are highly asymmetric and the
rachises curve posteriorly, as is characteristic of flight feathers
[15, 16]. The inner primaries (P1–P4) are straighter and more
symmetrical. Primary rachises are slender distally but thicker
proximally; the maximum rachis diameter of P4 is 1.8 mm.
Shallow grooves alternate with the primary feather shafts on
both left and right wings. Although previously interpreted as
a second set of impressions made by the wing, i.e., ‘‘double-
struck’’ impressions [7], these grooves are now recognized
as rachises lying dorsal to, and concealed by, the vanes of
the primaries [4, 6, 8, 9]. This is confirmed by close examina-
tion of the right wing, where the primaries are split away to
reveal rachises and barbs of these overlying feathers [4] (Fig-
ure 2), and also on the left wing, where a groove between P4
and P5 continues proximally to a calamus. Rachises of these
feathers angle outward relative to the primaries, as in dorsal
coverts [1].
Because they are comparable to the primaries in length, this
layer of feathers has been interpreted as dorsally displaced
primary remiges [4, 8]. However, this hypothesis is contra-
dicted by three lines of evidence. First, because their tips are
not visible at the wing margin, these feathers must be shorter
than the primaries. Second, the rachises are angled relative to
the primary rachises, instead of paralleling them as expected
for primaries. Third, interpreting these feathers as primaries
requires that every other primary has been displaced post-
mortem on both the left and right wings, without disturbance
of the other primaries or ventral coverts. No taphonomic
mechanism is known that could produce such a pattern. Inter-
preting these feathers as primaries undergoing a molt could
explain why they are shorter but does not explain their dorsal
position, the angling relative to the remiges, or the alternating
arrangement.
Instead, we propose an alternative explanation: the plumage
is preserved in life position, and the concealed feathers are
elongate dorsal coverts. The length, position, and orientation
of these feathers relative to the remiges all support this identi-
fication; in Neornithes, dorsal coverts are shortened relative to
the primary tract, lie dorsal to and alternating with the prima-
ries, and are angled outward relative to the primaries [1].
This interpretation is also corroborated by the London spec-
imen of Archaeopteryx, where the wing is preserved in dorsal
view, and several elongate feathers, identified as P1, S4, S7,
and S10 by de Beer [7], lie atop the primaries.
The ventral coverts are also visible. As in neornithines, the
ventral coverts of Archaeopteryx are angled inward relative
to the remiges; they are much longer than in most Neornithes,
however (Figure S2). Distally, the ventral coverts cover the
bases of P5–P8, but they cover most of P1–P4, and the
secondaries are almost completely obscured by the ventral
coverts, which extend nearly to the wing’s trailing edge. The
barbs of the ventral coverts do not interlock at their tips and
instead show the open pennaceous arrangement seen in
the ventral coverts of Neornithes. However, the barbs differ
from those of modern birds in being hyperelongate on the
trailing vane.
Wings of Archaeopteryx and Anchiornis
2263
The wing of Anchiornis huxleyi was studied and recon-
structed on the basis of BMNHC (Beijing Museum of Natural
History) PH828 (Figure 3), a partial skeleton preserving
feathers on both wings. Feathers (Figure 3) are preserved as
fossil melanosomes [17], with the dark tips of the remiges
being readily visible. Rachises and barbs are visible in places,
but elsewhere the overlapping feathers make it impossible to
discern details of feather structure. The tips of eight primaries
are visible on the left wing, but others appear to be concealed
by the secondaries, suggesting a higher primary count than in
Archaeopteryx. Primaries are straplike with tapered tips,
a feature present in some Anchiornis specimens [18] but
apparently not others [12]. Primaries are curved and symmet-
rical, with the tightly interlocking barbs that characterize
remiges [15, 19].
The primaries are overlain dorsally by a series of elongate
feathers, as in Archaeopteryx. Four tiers of feathers overlie
the primaries, which are approximately 90%, 75%, 50%, and
25% the length of the primaries (Figure 3). Barbs are tightly in-
terlocked, suggesting that these feathers are dorsal coverts.
An identical arrangement is seen in a second specimen [18]
(Figure S3).
On the left wing, the tips of five secondaries are visible, with
space for four to eight more. On the right, at least nine are
visible. Two rows of coverts, approximately 80% and 90%
the length of the secondaries, imbricate with the secondaries.
Secondaries and their coverts have narrow, symmetrical
Figure 1. Left Wing of Archaeopteryx lithograph-
ica, HMN 1880, Counterslab
(A) Left wing, ventral view.
(B) Line drawing of left wing.
(C) Closeup of primaries and dorsal primary
coverts.
(D) Line drawing of primaries and coverts.
Abbreviations: cov, shafts of major dorsal
primary coverts; P1–P8, primaries 1–8; S6–10,
secondaries 6–10.
vanes, as in the primaries, but have
rounded tips and straight rachises. In
contrast to Archaeopteryx and Avialae,
adjacent secondaries do not strongly
overlap one another, nor do adjacent
coverts show a strong overlap.
Discussion
The avian wing represents one of natural
selection’s most remarkable inventions.
Millions of years of evolution ultimately
modified the dinosaurian forelimb into
a highly efficient, feathered airfoil that
can rapidly change its span, shape,
and area. This design allows modern
birds to exploit a wide range of flight
kinematics and aerodynamic mecha-
nisms [20–23] and represents a key
innovation that has allowed dinosaurs
to rule the skies. Yet all flying Neornithes
are characterized by the same basic
wing configuration, in which long
remiges are overlapped at their bases
by short coverts [1–3]; living birds
therefore provide little evidence about the origins of this
wing design. Furthermore, if primitive birds [4–9] and feathered
dinosaurs had a similar configuration [10, 11], then this design
would appear to have been conserved for at least 150 million
years.
However, as shown here, Archaeopteryx and Anchiornis
depart markedly from modern birds in the arrangement of
the wing feathers, demonstrating that substantial evolution
occurred between early feathered dinosaurs and neornithines.
We propose the following scenario for the evolution of the
avian wing (Figure 4), using a recent phylogeny of Paraves
[24]. In winged dinosaurs, exemplified by Anchiornis, wing
feathers are unspecialized and undifferentiated (Figure 4).
Remiges are short, symmetrical, and straplike, with slender
rachises, and there is little differentiation between remiges
and coverts. Wing feathers are stacked into a series of
tiers, with coverts contributing substantially to the airfoil. In
Archaeopteryx, the feathers are more derived: the remiges,
especially the primaries, are elongated and have broader,
asymmetrical vanes. However, the shafts of the feathers are
still slender, and the wing retains long coverts. In the pygosty-
lian Confuciusornis sanctus (Figure S4), the wing is nearly
modern. Primaries are long and asymmetrical, and the coverts
are less than half the length of the primaries. The primaries also
have robust rachises, as in Neornithes. Assuming a mass of
180 g for Confuciusornis [25], an allometric equation derived
from modern birds [26] predicts a rachis diameter of 2.1 mm;
Current Biology Vol 22 No 23
2264
the actual diameter is 2.1–2.3 mm [27]. However, the wing
resembles Archaeopteryx in having a series of short primaries
forming the leading edge and in lacking an alula. A modern
wing, with elongate distal primaries and an alula, is first seen
in the Ornithothoraces, as exemplified by the enantiornithine
Eoalulavis hoyasi [28].
There are a number of issues that complicate this scenario.
First, the phylogenetic positions of Archaeopteryx and
Anchiornis remain uncertain. Most studies recover Archaeop-
teryx and Anchiornis as successive outgroups to modern
birds [12, 13, 24, 29, 30], but a recent analysis suggests that
Archaeopteryx and Anchiornis may be more closely related
to the Deinonychosauria [31]. If so, this would require that
either the long, asymmetrical remiges shared by Archaeop-
teryx and modern birds were acquired convergently, or that
the short and symmetrical remiges of Anchiornis are derived.
It could also mean that the multitiered feather arrangement
is derived for an Archaeopteryx-Deinonychosauria clade,
rather than primitive. Further complicating the scenario
presented here is the fact that the dromaeosaurid Microraptor
gui also has the long, asymmetrical primaries seen in Archae-
opteryx and Neornithes [11] (the morphology of the coverts,
however, remains unknown for Microraptor). Microraptor
may have independently evolved the advanced feather
morphology, or again, Anchiornis may represent a reversal. It
is also possible that the derived morphology of the remiges
seen in Archaeopteryx and Microraptor is primitive for Para-
ves, and that Anchiornis actually lies outside of this clade. To
resolve these conflicts, we require both a better understanding
of maniraptoran phylogeny and new information on the
plumage of basal maniraptorans such as Protarchaeopteryx
and Caudipteryx [32].
Regardless of the precise scenario invoked, it is clear that
Archaeopteryx and nonavian dinosaurs have a wing feather
organization that differs from that of modern birds. An
advanced wing morphology did appear relatively early in avian
evolution, however. The existence of a modern wing in the
Enantiornithes indicates that this morphology predates the
enantiornithine-ornithurine split. The oldest known enantiorni-
thine is Protopteryx fengningensis [33], which comes from
strata dated to 131 Ma [34], roughly 25 million years after
Figure 2. Right Wing of Archaeopteryx lithog-
raphica, HMN 1880, Counterslab
(A) Right wing, ventral view.
(B) Line drawing of right wing.
(C) Closeup of primaries and coverts.
(D) Line drawing of primaries and coverts.
Abbreviations: cov, major dorsal primary coverts;
P1–P8, primaries 1–8; S8–10, secondaries 8-10;
VC, major ventral coverts.
Anchiornis, which occurs in rocks dated
to 155 Ma [12]. Thus, the wing feather
arrangement seen in modern birds may
have evolved within a period spanning
perhaps a few tens of millions of years
and then remained largely unchanged
for over 130 million years. This punctu-
ated pattern is also seen in Pterosauria,
which retain a similar wing configuration
from the Triassic to the end of the Creta-
ceous, a period of 140 million years or
more [35], and in Chiroptera, which
evolved a modern wing morphology within 15 million years of
the extinction of the dinosaurs and retained this design for
more than 50 million years [36]. Strikingly, a pattern of stasis
is found in manmade aircraft as well: following rapid advances
in aircraft design in the early 20th century, progress slowed in
later decades, such that many aircraft designed in the mid-
20th century still operate. The processes behind the evolution
of vertebrate wings and aircraft wings may be the same. The
constraints imposed by fluid mechanics mean that a relatively
small number of possible wing configurations are effective
airfoils [37]; once these geometries are discovered either by
natural selection or aeronautical engineering, only small refine-
ments are possible.
The wing feather arrangement described here also has
important implications for understanding the flight ability of
Archaeopteryx and Anchiornis. It has recently been argued
that the feather shafts of Archaeopteryx are more slender
than expected for a bird of its size, which may have limited
the ability of the feathers to function as an airfoil [14].
However, there are several issues with this analysis. First,
the study used an incorrect mass estimate for Archaeop-
teryx; a mass of 276 g is assumed for the Munich Archaeop-
teryx [14], yet the reference cited [9] estimates its mass as
222 g, and the animal may have weighed as little as 150 g
[25]. Second, the diameter of the primary feather shafts
was underestimated [25] (as confirmed by our examination
of casts and photographs of the Munich specimen). Using
masses of 252 g for the Berlin specimen [38] and 222 g for
the Munich specimen [9], the allometric equation calculated
from modern birds [26] predicts a primary feather shaft diam-
eter of 2.4 mm for the Berlin specimen and 2.3 mm for the
Munich specimen. Measured rachis diameters in the Berlin
and Munich specimens are 1.8 mm and 1.4 mm—75% and
61% of the predicted values, respectively. Using a lower
mass estimate of 150 g for the Munich Archaeopteryx [25],
the observed diameter is still 71% of the predicted diameter
of 2.0 mm. Therefore, rachis diameter of Archaeopteryx is
lower than expected for an extant bird of comparable size,
although the difference is less than previously reported [25].
Anchiornis has also been described as having slender feather
shafts [12], and although coverts cover the bases of the
Wings of Archaeopteryx and Anchiornis
2265

Figure 3. Wing of Anchiornis huxleyi BMNHC PH828

(A) Left wing.
(B) Line drawing of left wing.
(C) Detail of outer wing.
(D) Line drawing of outer wing.
(E) Detail of inner wing.
(F) Line drawing of inner wing.
Abbreviations: P, primaries; PC1, dorsal major primary coverts; PC2, dorsal median primary coverts; PC3 and PC4, dorsal minor primary coverts; S, second-
aries; SC1, major secondary coverts; SC2, median secondary coverts.

rachises in BMNHC PH828, the distal rachis does appear to
be relatively slender.
The slender primary feather shafts in Archaeopteryx and
Anchiornis therefore raise questions about the ability of these
wings to support their owners in flight. The feather arrange-
ment described above, however, may provide an answer. By
packing together many layers of relatively weak feathers, the
wings of Archaeopteryx and Anchiornis may have been strong
enough to function as airfoils. The multiple layers of feathers
would also have resulted in an airfoil that was much thicker
than that of modern birds. Whereas this arrangement would
have increased wing profile drag at low speeds, at higher
speeds the thick wing section would decrease profile drag [37].
Although the wings of Archaeopteryx and Anchiornis
may have been efficient as airfoils, their primitive feather
configurations may have prevented their wings from
functioning in the same fashion as those of Neornithes. First,
many extant birds can spread the tips of the primaries apart
to create a slotted wingtip. This arrangement decreases lift-
induced drag by spreading wingtip vortices vertically [39],
and these slots are particularly important in slow flight,
because induced drag is strongest at low speeds [21, 37].
The long coverts in Archaeopteryx and Anchiornis would
have covered gaps between the primaries, however, prevent-
ing the wingtip from assuming a slotted configuration. Second,
modern birds can separate the flight feathers on the upstroke
to assume a louvered arrangement, by rotating the feathers
about their long axes [16]. This ‘‘Venetian blind’’ mechanism
creates openings between the primaries that allow air to
pass through [40] and is used by many birds in low-speed flap-
ping flight [40], hovering [16], and wing-assisted incline
running [41] to permit a rapid upstroke. In Archaeopteryx and
Current Biology Vol 22 No 23
2266
Anchiornis, however, the long coverts would have spanned the
gaps between the primaries, preventing the wing from
assuming a louvered configuration at low speeds.
If Archaeopteryx and Anchiornis lacked the wing mor-
phology seen in extant avians, then we cannot automatically
assume [41, 42] that their wings functioned like those of living
birds. Instead, evolution of the form of the wing suggests that
the function of the wing has evolved as well. In particular, the
inability of the wings of Archaeopteryx and Anchiornis to
employ slots or feather separation suggests that the ability
to take off or fly at low speeds may have been limited, and
that instead the wings functioned primarily in high-speed
gliding or flapping flight. This, in turn, would lend support to
a trees-down origin of avian flight, in which the evolution of
powered flight was preceded by arboreal parachuting and
gliding [21, 43].
Experimental Procedures
The wing of Archaeopteryx was imaged from the counterslab of HMN 1880
by using a Canon XSI DSLR and Tamron 90 mm macro lens to take an
exposure-bracketed set of three images, which were then combined in
Adobe Photoshop to create a high dynamic range (HDR) image, improving
contrast of individual barbs in highlighted and shadowed parts of the
fossil. Anchiornis BMNHC PH828 was imaged using a Nikon d90 DSLR
and Nikkor 60 mm micro lens. Overall image contrast and color channels
were adjusted in Adobe Photoshop, and illustrations were made by tracing
photographs in Adobe Illustrator. Predicted feather shaft diameters for
Archaeopteryx and Confuciusornis were calculated using the equation
diameter = 0.004(mass)0.37 [26].
Supplemental Information
Supplemental Information includes four figures and can be found with this
article online at http://dx.doi.org/10.1016/j.cub.2012.09.052.
Acknowledgments
We thank Daniela Schwarz-Wings for access to the Berlin Archaeopteryx.
N.R.L. was supported by the Yale Institute for Biospheric Studies. We
Figure 4. Hypothesized Evolution of the Avian
Wing
(A) Ancestral wing morphology (exemplified by
Anchiornis), where the wing is composed of
slender, symmetrical, and poorly differentiated
flight feathers.
(B) Wing of early Avialae (exemplified by Archae-
opteryx), where the remiges are elongated,
broad, and asymmetrical.
(C) Wing of pygostylia (here Confuciusornis), in
which the primary remiges are further elongated
and the coverts are shortened. Phylogeny after
Turner et al. [24].
also thank Julia Clarke, Daniel Field, Mark Norell, and Rick Prum for
discussions.
Received: August 18, 2012
Revised: September 21, 2012
Accepted: September 24, 2012
Published: November 21, 2012
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