Journal of Zoology

Journal of Zoology. Print ISSN 0952-8369

REVIEW

The evolution of ‘bizarre structures’ in dinosaurs:

biomechanics, sexual selection, social selection or
species recognition?
K. Padian1 & J. R. Horner2
1 Department of Integrative Biology and Museum of Paleontology, University of California, Berkeley, CA, USA
2 Museum of the Rockies, Montana State University, Bozeman, MT, USA

Keywords
Dinosauria; functional morphology; sexual
selection; social selection; species
recognition.
Correspondence
Kevin Padian, Department of Integrative
Biology and Museum of Paleontology,
University of California, Berkeley, CA, USA.
Email: kpadian@berkeley.edu
Editor: Steven Le Comber
Received 14 October 2009; revised 4
February 2010; accepted 22 April 2010
doi:10.1111/j.1469-7998.2010.00719.x
Abstract
‘Bizarre structures’ in dinosaurs have four main traditional explanations: mechan-
ical function, sexual selection, social selection and species recognition. Any of
these can be plausible for individual species, but they fail to be persuasive when
other lines of evidence cannot adequately test them. The first three also fail as
general propositions when phylogenetic analyses based on other characters do not
support scenarios of selective improvement of such functions in their clade (or the
explanation simply does not apply to any other species in the clade). Moreover, the
hypothesis of sexual selection requires significant sexual dimorphism, which has
never been conclusively established in dinosaurs.
We propose instead that species recognition may have been a more general
force that drove the evolution of bizarre structures in dinosaurs. That is, the
bizarre structures communicate to other individuals a variety of possible associa-
tional cues, including species identification, potential protection and social habits
and the appropriateness of potential mates. In other words, bizarre structures
amount to an advertisement for positive association. Neither species recognition
nor any other hypothesis should be a ‘default’ explanation. Although direct
observation is impossible, we propose two tests. First, contrary to adaptive, social
or sexual selection, under the species recognition model morphology should be
expected to evolve without obvious directional trends, because the only objective is
to differ from one’s relatives. Hence, patterns of evolution of bizarre structures
should be relatively proliferative and non-directional. Second, several contem-
poraneous species should overlap in geographic range (sympatric, parapatric,
peripatric). Fossil species often show evidence of this pattern in the past by ‘ghost
ranges’ of related taxa. These tests together could reinforce or weaken an argument
for species recognition.

argue for a particular theory that explains everything than

Introduction
‘Bizarre structures’ in dinosaurs and other extinct animals
(e.g. Gould, 1974) are of perennial interest to paleontolo-
gists and have become a staple of textbooks on evolution
because they raise perennial questions. What did these
structures do? How did they evolve? If they were so useful,
how did they contribute to their bearers’ evolutionary
success? If their bearers are extinct, did they become a
liability at some point?
In this paper, we explore the principal explanations for
the evolution of ‘bizarre structures.’ The kinds of explana-
tions we discuss include the teleology of what they were for
and how they evolved. We recast these explanations using
current methods of comparative biology. Our goal is less to
to suggest how these kinds of evolutionary problems should
be addressed, and to suggest some criteria for testing them.
Our hope is that others will both improve on our suggestions
and bring new data to the questions.
By ‘bizarre structures’ we mean features that are unusual
enough, to the trained eyes of paleobiologists, to invite
explanations beyond the basic functions of feeding, locomo-
tion, respiration and so on (Farlow & Dodson, 1974; Gould,
1974; Molnar, 1977; Main et al., 2005). In many respects
these structures are similar (but not necessarily analogous)
to certain structures in living animals. They include the frills
and horns of ceratopsians, the domes of pachycephalosaurs,
the crests of lambeosaurine hadrosaurs, the scute complexes
of ankylosaurs and the plates and spikes of stegosaurs. We

Journal of Zoology 283 (2011) 3–17

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‘Bizarre structures’ in dinosaurs
discuss four general types of explanations: mechanical func-
tion, sexual selection, social selection and species recogni-
tion. The first two of these are pre-eminent in
paleobiological explanation (e.g. Galton, 1970; Farlow &
Dodson, 1974; Dodson, 1975; Hopson, 1975; Farlow,
Thompson & Rosner, 1976; Molnar, 1977; Buffrenil, Far-
low & de Ricqlès, 1986; etc.). The third has been advocated
most recently and thoroughly by Hieronymus et al. (2009).
The fourth has not been extensively considered by any
authors, although it has been frequently acknowledged in
functional and behavioral considerations (e.g. Farlow &
Dodson, 1974; Hopson, 1975; Molnar, 1977; Sampson,
1999; Hieronymus et al., 2009). There has been an historical
predilection to attempt first to explain a bizarre structure in
mechanical terms; if this explanation appears weak or is
contraindicated, it has been traditional to attribute the
feature to ‘sexual display’ by virtue of its apparent useless-
ness for mechanical function. In this way, sexual display has
often become a ‘default’ explanation that was seldom
explicitly tested or questioned.
We acknowledge several classes of facts. First, some
structures may have served more than one function. For
example, ankylosaur armor may have been defensive but
also distinctive enough to have served a role in species
recognition. After all, exaptation is a pre-eminent factor in
macroevolutionary change. Second, because many soft part
features and also nearly all behaviors are not preserved in
the fossil record, affairs may have been far more complex
than paleontologists can detect. We also allow that not
enough is known to determine the origin of some features;
for example, there are too few known specimens of cranially
adorned theropod taxa such as Dilophosaurus, Cryolopho-
saurus and Carnotaurus to permit a test of evolutionary
explanations. We see no reason to be dogmatic about
particular hypotheses, and no reason not to be pluralistic
about explanations when appropriate. Our goal is to pro-
pose a set of explicit tests of mechanical and behavioral
hypotheses that we hope will set up discriminatory criteria
for these kinds of explanations.
4 Journal of Zoology 283 (2011) 3–17
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K. Padian and J. R. Horner
Methods
Although there are many approaches to explaining mor-
phology in extinct organisms (Hickman, 1980), inferences
about function and behavior are based on two general
models: homology and analogy (essentially, historical and
ahistorical explanations: Weishampel, 1997). The accepted
approach to evaluating homology of function and behavior
in extinct animals is Witmer’s (1995) extant phylogenetic
bracket (EPB). For this purpose, a phylogeny of living and
related fossil forms is required. The degree to which a
condition can be inferred reliably as present in an extinct
taxon is related to its position among living forms that are
known to share the function or behavior (Fig. 1). Because
crocodiles and birds, the two extant brackets of extinct
dinosaurs, share none of the bizarre structures of extinct
dinosaurs, the EPB cannot provide much direct guidance on
these problems. There are simply no available homologous
structures, with the possible exceptions of the cranial crests
of lambeosaurine hadrosaurs and cassowaries, and the
scutes of crocodiles and thyreophorans (which, being absent
in their respective common ancestors, must be regarded as
parallelisms, despite an obvious homological basis in bone
histology: Scheyer & Sander, 2004; Main et al., 2005).
Analogy to living forms is the approach that remains
when arguments of homology cannot be made, and it is even
more problematic. The quality of an explanation depends in
part on the precision of definition of the features that are
compared, and the separation of those features (and func-
tions) from ancillary or irrelevant ones (Whewell, 1859;
Padian, 1995; Wilson, 1998).
Classes of explanation of
bizarre structures
The two general classes of explanation of bizarre structures
in dinosaurs relate to function and display (including sexual
selection, social selection and species recognition). Each
kind of explanation has a long history in the literature,
Figure 1 The Extant Phylogenetic Bracket (Wit-
mer, 1995) of extinct dinosaur clades that
characteristically and independently evolved
‘bizarre structures:’ stegosaurs and ankylo-
saurs (parallel elaboration of homologously
shared dermal scutes), lambeosaurine hadro-
saurs, neoceratopsians, pachycephalosaurs
and ceratosaurian theropods. The bizarre
morphologies of these dinosaurian groups are
not shared by their living bracket, crocodiles
and birds; crocodiles have scutes, but they are
never elaborated like those of thyreophoran
dinosaurs.
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K. Padian and J. R. Horner
Table 1 Some proposed functions of ‘bizarre’ structures
Mechanical
Defense
Communication
Thermoregulation
Sensory
Procuring food
Display
Interspecific
Deter predators
Discourage association of non-conspecifics
Intraspecific
Intrasexual
Establish territory
Ward off rivals for resources (including mates)
Encourage association of conspecifics
Compete for resources
Intersexual
Attract mates
Encourage association of conspecifics
Note that functions of species recognition encompass interactions
both between species (discourage association of non-conspecifics)
and within species (‘encourage association of conspecifics,’ both of
the same sex and of different sexes).
including discussions of dinosaur behavior (Horner & Gor-
man, 1988; Carpenter, Hirsch & Horner, 1994; Currie &
Padian, 1997; Farlow & Brett-Surman, 1997; Horner &
Dobb, 1997; Carpenter, 1999; Weishampel, Dodson &
Osmolska, 2004; Hieronymus et al., 2009). We summarize
these classes of explanation in Table 1.
It is important that we define our terms. Mechanical
function refers to a specific adaptation such as feeding,
locomotion, insulation or communication. Sexual selection
is the advantage gained to access to mates when one
sex possesses a specific feature that the other does not,
and uses it to attract mates or repel rivals for mates (Darwin,
1859, 1871). We want to emphasize here the importance
of a discrete structure, function or behavior present in
one sex but not the other, that is used for these two
purposes. We also emphasize that this true sexual dimorph-
ism is different from a simple ‘sexual difference’ in
which one sex is slightly larger or more robust than the
other, but possesses no particular structures for these
purposes. (We recognize that there is debate about this
among behavioral ecologists, and we discuss it elsewhere.)
Social selection refers to features that individuals in a
species use to improve their competitive advantage for
resources. Species recognition refers to features that allow
others of the same species to recognize each other for
various social purposes. Mate recognition is not the same
thing, but it is a subset because it is important for individuals
to mate with others in the same species. We want to state
emphatically that we do not reject the possible operation of
any and all of these processes in extinct dinosaurs in
principle. We ask how well established any and all of these
are in specific cases.
Journal of Zoology 283 (2011) 3–17
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‘Bizarre structures’ in dinosaurs
Mechanical explanations
Explanations of individual taxa
Many possible mechanical explanations have been proposed
and tested for various bizarre skeletal features of individual
dinosaur species (Weishampel, 1981, 1997; Farke, 2004;
Farke, Wolff & Tanke, 2009; Hieronymus et al., 2009). In
our view, Weishampel’s (1981) classic study of the crest of
the hadrosaur Parasaurolophus is a model for examining
functional inferences in extinct individual taxa. Weishampel
first divided all proposed hypotheses into testable and
untestable, and then proceeded to see if the testable ones
could be falsified or supported by other lines of evidence. He
found that most hypotheses of display and behavior could
not be explicitly tested, but some mechanical functions, such
as snorkeling, head-butting and air storage, could be tested
and rejected. Weishampel tested the proposed function of a
resonance chamber by building a model of the nasal pas-
sages and diverticula, and passing a spectrum of oscillating
frequencies through them. Certain frequencies, as expected,
resonated better than others, and Weishampel indepen-
dently tested this outcome by determining whether the
auditory organs were well attuned to those frequencies by
studying the size and morphology of the stapedial region.
Whereas this study did not ‘prove’ any particular function,
and could not logically rule out several weakly supported or
untestable explanations (see Weishampel, 1997), it is a
model study for testing functional hypotheses of individual
organisms in paleobiology.
But Weishampel’s approach, thorough as it was, did not
account for all aspects of the problem, as he recognized. He
noted one: the characters related to vocalizing and hearing
reflect different (if not contradictory) respective phyloge-
netic brackets (Weishampel, 1997). In other words, other
character state distributions do not match, so they did not
apparently evolve in step. This is a specific problem for that
case. Disjunct sets of character distributions cannot support
a unified functional hypothesis that purports to explain the
evolution of an adaptation (although in this case an exapta-
tion may be possible).
One shortcoming of most functional explanations for
bizarre structures in extinct dinosaurs is that the evolution
of these features and functions in a clade is very seldom
considered. Without doing so, there is no evidence that the
function (in the sense of an adaptation) evolved at all, and
therefore the hypothesized function itself must be consid-
ered in doubt, unless there is good independent evidence of
it. The demonstration of its evolution requires a phyloge-
netic component.
Phylogenetic dissection of adaptation (PDA)
When paleobiologists discuss functions of bizarre struc-
tures, they are generally discussing adaptations. It is a
truism of evolutionary biology that adaptations are shaped
by natural selection (Williams, 1992). Paleobiologists can-
not assess selection in populations through generations, as
5
‘Bizarre structures’ in dinosaurs
microevolutionists can (e.g. Endler, 1986; Brandon, 1996).
But they can assess natural selection at a more general
hierarchical level in lineages, living and extinct, by mapping
the elaboration of structures and the improvement of
proposed functions upon phylogenies based on other char-
acters (e.g. Padian, 2001; Padian & Horner, 2002, 2004).
In order for an adaptation to be assessed (Padian, 1982,
1987), its necessary components must be identified and
separated from non-essential ones. By plotting these char-
acter states on a phylogeny built from other characters, the
assembly of the adaptation can be traced. Even after the
basic adaptation is assembled, further modifications can be
tracked in the same way (Padian, 2001). This method of
PDA can be formalized in the following way (modified from
Padian, 1982, 1987, 1995, 2001):
1. Identify the adaptation, its diagnostic (vs. merely asso-
ciated) features and the groups that possess it.
2. Perform phylogenetic analyses of the groups, including
closest sister taxa, using all available character taxa.
3. Identify the phylogenetic sequence of acquisition of each
diagnostic feature of the adaptation.
4. Analyze the apparent roles, if any, of diagnostic char-
acters at each successive stage before the adaptation is
assembled, using functional, physical, ecological, genetic
and other lines of evidence.
The implication of this method for the assessment of
bizarre structures in dinosaurs is that, if such explanations
are to move beyond the ad hoc, they must be able to explain
the evolution of these features, the assembly of their char-
acters and functions. In other words, at successive nodes
along the spine of the cladogram, one should be able to
point to specific characters diagnostic of the proposed
adaptation, and assess their function with respect to the
organism as a whole. Such assessments need to take into
account the roles of other features in the functional complex
in order to provide an adequate cross-test (Padian, 2001).
Moving to successive nodes along the spine of the clado-
gram, the evolution of the features from stage to stage
should emerge. If there is no evidence for the improvement
of a function or the assembly of a new one, the adaptive
hypothesis fails. Therefore, functional explanations that are
not tested phylogenetically have no demonstrated evolu-
tionary basis and are of limited value (Fig. 2; Weishampel,
1997).
Testing mechanical hypotheses
We divide these into four general (and not mutually exclu-
sive) classes: defense, communication, thermoregulation
and sensory function.
Defense
These features can be attributed to repulsion of predators
and to conspecifics of the same sex in agonistic behaviors
(non-exclusively). Notable examples are the horns and frills
of ceratopsians, the plates and spikes of stegosaurs, the
scutes and tail club of ankylosaurs and the domes of
6 Journal of Zoology 283 (2011) 3–17
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K. Padian and J. R. Horner
Figure 2 Phylogeny as a test of the assembly of an adaptation.
Consider an adaptation defined by five necessary features (1–5)
following a PDA analysis (Padian, 1995, 2001; see text). Phylogenetic
analysis reveals the clade that shares this function (and thereby
shares all five features); it also reveals the order of assembly of those
five features on nodes along the spine of a more inclusive cladogram.
At each of the nodes along this spine (which have successive subsets
of those five features), a different function can be proposed and
tested; new secondary (exaptive) functions and improvements in
existing functions may also be proposed and tested.
pachycephalosaurs. Weishampel (1981) tested the possibi-
lity of a defensive function of lambeosaurine crests and
concluded that the bone was too thin to have been of any
use in this regard.
Ankylosaurs would seem to pose the least controversial
example of a defensive function for bizarre structures, in this
case the dermal scutes (traditionally and tellingly called
‘armor’) and tail ‘club’ (in ankylosaurids only: Carpenter,
1997, 2001; Vickaryous, Maryanska & Weishampel, 2004).
Scutes cover the skull, the neck, the back, and much of the
tail, but there is great variety in their size, form and extent
among ankylosaurs (Carpenter, 1997). This suggests that
there was no ‘optimal’ pattern of scute form and distribu-
tion, and therefore it is difficult to propose that a defensive
function was successively ‘improved’ in ankylosaurs. How-
ever, consideration of their outgroups shows that ankylo-
saurs had more extensive dermal ossifications than the basal
thyreophorans Scutellosaurus and Scelidosaurus (the latter
often considered an ankylosaur), not to mention the stego-
saurs, which lost all but the parasagittal rows (Main et al.,
2005). This pattern points to defense as a plausible basal
function of ankylosaur scutes, and suggests that whatever
the variations in scute form and distribution, they were
‘good enough’ to serve an adequate defensive function.
Yet, as Carpenter (1997: p. 315, fig. 22.6) notes, the varia-
tion in scute form, and notably in the more conspicuous
long neck spikes, suggests no obvious defensive strategy (see
also Scheyer & Sander, 2004), and may instead be primarily
related to display. Sexual dimorphism has not been estab-
lished, so sexual selection has no support, but social selec-
tion (Hieronymus et al., 2009) could be investigated further.
Several evolutionary strategies may have been involved
here. The enlarged and fused scutes at the end of the
ankylosaurid tail, preceded by a series of fused caudal
vertebrae, have often been invoked as a weapon, and this
seems to be supported by the enlarged areas of muscle
attachment on the pelvis, hindlimbs and transverse
c 2010 The Zoological Society of London
K. Padian and J. R. Horner
Figure 3 Cladogram of advanced ceratopsians, after Dodson et al.
(2004). No specific functional or behavioral hypotheses are currently
proposed (or have been tested) to account for evolutionary trends in
the features related to the bizarre cranial characters of these taxa.
processes of the anterior caudal vertebrae, despite some
limits in vertical mobility (Vickaryous et al., 2004).
Most attributions of defense to the frills of neoceratop-
sians have focused on Triceratops (Fig. 3). This is apparently
because Triceratops has prominent orbital horns as well as a
solid frill, so its function in ‘jousting’ is easily visualized (e.g.
Farke, 2004). However, Triceratops is virtually (along with
Avaceratops) the only neoceratopsian with a solid frill,
which is also the shortest among large neoceratopsians
(Fig. 3). Other large neoceratopsians have substantial open-
ings in their frills, which would have been of little use in
defense. It now turns out that the adult Triceratops is in fact
what has been called Torosaurus, and its frill is not only
fenestrated but also quite thin, as in other neoceratopsians
(Scannella & Horner, in press). We hypothesize that,
Journal of Zoology 283 (2011) 3–17
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‘Bizarre structures’ in dinosaurs
because it is so similar to young Triceratops, the adult form
of Avaceratops may turn out to have been fenestrated as
well.
And horns vary widely; chasmosaurines had orbital horns
of various sizes and orientations, but most centrosaurines
had small orbital horns, and nasal horns of variable size that
show no obvious function in combat (Farke et al., 2009).
Farke (2004) used restored scale models of Triceratops to
determine how individuals might have fought each other,
interlocking horns, and Farke et al. (2009) showed that
injuries occurred significantly more often on skull bones that
would have been expected according to his predictions.
However, even if this function is plausible, it has not been
proposed and tested for other chasmosaurines, although it
was absent in centrosaurines (Farke et al., 2009). The most
recent published phylogenies of neoceratopsians (Xu et al.,
2002; Dodson, Forster & Sampson, 2004; Fig. 3) show no
directional pattern of improvement of either brow horns or
nose horns. Hence there is no evidence for adaptation to a
particular function, and other hypotheses also need to be
considered as a general explanation for the evolution of
horns and frills.
For stegosaurs, as Main et al. (2005) have shown, the
elaboration of plates and spikes shows no phylogenetic
trends in adaptation to proposed functions of thermoregu-
lation (Galton & Upchurch, 2004b). The possible function
of defense has been rejected by several authors (Buffrenil
et al., 1986; Main et al., 2005): the plates consist of a thin
layer of compact bone surrounding a central core of well-
vascularized, lattice-like (spongy) trabecular bone that
would be crushed easily by the teeth of any large theropod.
A possible function in deterring predators by making the
animal appear larger has been suggested, but again it would
not explain why Stegosaurus has large plates and those of
the contemporaneous Kentrurosaurus and others are much
smaller.
Pachycephalosaur domes have been assumed to have
been used in head-butting, ever since Colbert’s (1955) casual
suggestion (review in Maryanska, Chapman & Weishampel,
2004). However, histological studies have shown that the
columnar cell structure of these domes would not have
deflected the forces incurred in battering, as reasonably
proposed by Sues (1978) on the basis of biomechanical
models of gross anatomy. The spongy bone that was
thought to be protective of the brain during head-butting,
by analogy to similar bone in bighorn sheep (Galton, 1970),
is actually characteristic of juvenile skulls; the skulls of
adults, in which most head-butting would have been ex-
pected to occur, have compact bone in their external cortices
(Goodwin & Horner, 2004). Moreover, the spongy bone of
juvenile skulls is organized in such a way that reflects radial
growth of the bone, which indicates rapid growth (Francil-
lon-Vieillot et al., 1990: p. 512). Rather than deflecting
concussive forces from the brain cavity, this radial organiza-
tion would have more likely directed them into the brain
cavity (Goodwin & Horner, 2004).
Maryanska et al. (2004) recently renewed the argument
for mechanical agonistic behavior, but their analysis had no
7
‘Bizarre structures’ in dinosaurs
control for ontogeny or sexual dimorphism, so there is no
support for assigning male status to larger and thicker
domes as they did. Moreover, the knobs and spikes that
ornamented some pachycephalosaur skulls (such as Stygi-
moloch) would not have been visible until the heads were
lowered, and in any case could not have been involved in
combat (Goodwin, Rosner & Johnson, 1998). For these
reasons a function in combat for both the domes and
ornamentation is implausible. Moreover, there is now evi-
dence that Stygimoloch was a subadult form of Pachycepha-
losaurus, which has somewhat less extreme spikes than
Stygimoloch, thus casting doubt on the functional interpre-
tation (Horner & Goodwin, 2009).
Communication
Weishampel’s (1981, 1997) study of Parasaurolophus de-
scribed above was the first example of an explicit test of a
hypothesis that a particular structure functioned in commu-
nication. As noted, this function may apply to this genus,
but it has not been proposed and tested for other lambeo-
saurines until recently, when Evans, Ridgely & Witmer
(2009) examined Lambeosaurus, Corythosaurus and Hypa-
crosaurus. They showed, as Weishampel (1981) had done
using Lophorhothon, that the ear region was capable of
hearing the low-frequency sounds that Weishampel calcu-
lated might have been produced by the resonating crests of
these hadrosaurs. However, phylogenetic analysis of lam-
beosaurines (Horner, Weishampel & Forster, 2004) shows
no apparent trends in selection for improvement of the
features related to this function (Fig. 4), and Evans et al.
Corythosaurus
Hypacrosaurus
Lambeosaurus
Parasaurolophus
Tsintaosaurus
Figure 4 Cladogram of lambeosaurine hadrosaurs, after Horner et al.
(2004). No specific functional or behavioral hypotheses are currently
proposed (or have been tested) to account for evolutionary trends in
the features related to the bizarre cranial characters of these taxa.
8 Journal of Zoology 283 (2011) 3–17
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K. Padian and J. R. Horner
(2009) noted that Hypacrosaurus altispinus had a particu-
larly derived and convoluted nasal chamber.
Thermoregulation
Only two kinds of dinosaurian structures have been pro-
posed as thermoregulatory structures. The first is the plates
of stegosaurs. Main et al. (2005) showed that the explana-
tion hypothesized for stegosaurs (Buffrenil et al., 1986)
could not be completely eliminated for Stegosaurus itself
but was unlikely to apply to related taxa, so there was no
evidence of the evolution of a functional adaptation in the
group. The other example is the frills of ceratopsians; like
the plates of stegosaurs, these structures bear numerous
superficial vascular grooves that could be interpreted as
conductors of blood vessels that could modify body core
temperatures (Rigby Jr, 1990). However, this hypothesis has
never been rigorously tested, despite some intriguing evi-
dence (Barrick et al., 1998), and it is more conservative to
suppose that the blood vessels nourished the rapid growth of
frills and plates, which seem to have become more elabo-
rated at the sub-adult stage (Horner & Marshall, 2002;
Dodson et al., 2004; Main et al., 2005).
Sensory
Ostrom (1961, 1962) proposed that the crest of Parasaur-
olophus-enhanced olfaction: that is, an extended nasal
epithelium with sensory cells may have improved the ani-
mal’s ability to smell. However, as Hopson (1975) noted,
lambeosaurine crest variability is too great to be explained
simply by selection for olfaction. Moreover, lambeosaurines
had no particularly specialized or enlarged olfactory lobes in
the brain, compared with other dinosaurs (Ostrom, 1961;
Evans et al., 2009).
Procuring food
Bizarre structures such as tusks are used by some animals to
procure food, but to our knowledge no such function has
been seriously proposed or tested for dinosaurs.
Display
Display functions can be divided broadly into antagonistic
versus attractive: the repulsion of various threats versus the
attraction of potential mates (Table 1). But sometimes, as in
many mammals and some birds, these functions are related
(Darwin, 1871). Attraction only applies to the other sex of
the same species, but not all structures involved here fall into
the category of sexual selection.
Interspecific
Hypotheses about structures that may play a role in repel-
ling potential predators are difficult to test. Buffrenil et al.
(1986) determined that the plates of stegosaurs were not well
constructed to resist the bites of predators such as Allo-
saurus. The plates may have made the animals look larger,
c 2010 The Zoological Society of London
K. Padian and J. R. Horner
and this function may also be attributed to most bizarre
cranial structures of dinosaurs, as well as to the plates of
ankylosaurs (Carpenter, 1997). However, it is difficult to
know how to test this hypothesis. Moreover, the evolution-
ary literature suggests that structures hypothesized to repel
predators in living forms, whether by aposematic mimicry or
agonistic display, do not appear to enjoy long-term success
unless the threat they promise can be fulfilled (Futuyma,
2009).
Intraspecific
(i) Intrasexual: Females seldom contest each other, except to
establish social hierarchies (as in some mammals that travel
in social groups or herds), but males commonly contest
males, among both invertebrates (notably arthropods) and
vertebrates (Darwin, 1871). In general, territory and re-
sources form the basis of male competition in mammals
and in birds. Possession of resources is usually linked to
competitive superiority among males, and this advantage in
turn makes males more able to secure females, or more
attractive to females, because females are thought to per-
ceive greater advantage in mating with these males. (Some
birds short-circuit the process or use a proxy to attract
females through colorful feathers or eloquent songs [Dar-
win, 1871; Andersson, 1994].) Some bizarre structures in
extinct dinosaurs may have threatened rivals, but this is
difficult to test without direct knowledge of behaviors that
are not preserved in the fossil record.
(ii) Intersexual: The principal means of intersexual dis-
play is display for mates, traditionally called sexual display.
Sexual display usually implies sexual selection, and explana-
tions of sexual selection must be evaluated much like those
for mechanical adaptations. In contemporary populations,
sexual selection often acts on minor features and elaborates
them (Mendelson & Shaw, 2005); intense sexual selection
can result in runaway selection (Futuyma, 2009) and (or)
divergent selection (Kroodsma et al., 1985; Price, 1998).
Evolutionary theory holds that this kind of divergence can
result in speciation (Futuyma, 2009), and that like natural
selection, sexual selection can be responsible for patterns of
sorting in clades (Vrba, 1984; Sampson, 1999). This could be
shown if the characters subject to sexual selection show non-
random trends in clades (though the variation of the trends
themselves does not have to be directional or trendlike).
A problem with invoking sexual display as the explana-
tion of bizarre structures can be traced to Darwin’s (1871)
original formation of the problem of sexual selection.
Darwin emphasized that sexual selection could only apply
when one sex bears structures used in intersexual display (or
agonistic behavior in intrasexual interaction). In other
words, sexual selection cannot be invoked without discrete,
qualitative features of sexual dimorphism. (We acknowledge
that many neobiologists [apparently originating with West-
Eberhard’s, 1983 conflation of the concepts] feel that sexual
dimorphism is not necessary for sexual selection, but Dar-
win defined the concept in this way and by definition he
cannot be wrong. This does not deny that various other
Journal of Zoology 283 (2011) 3–17
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c 2010 The Zoological Society of London
‘Bizarre structures’ in dinosaurs
phenomena associated with competition for mates and
reproductive success are interesting and important; but they
are not strictly part of sexual selection.) Unfortunately, this
degree of sexual dimorphism, typical of birds and some
mammals, has not been sufficiently established for dino-
saurs.
(iii) Social selection: This concept (West-Eberhard, 1983)
was recently applied to dinosaurs by Hieronymus et al.
(2009), who argued persuasively that the nasal cornifications
of centrosaurine ceratopsians were progressively selected for
larger size and broader display. According to them, ‘social
selection occurs when there is differential success in within-
species competition for any limited resource.’ Two problems
with this definition, as applied to fossils, are that within-
species phenomena can almost never be observed, and
competition is particularly difficult to establish in extinct
forms (Benton, 1996). On the other hand, it is possible to
identify structures that can plausibly have functioned only
in social interaction (as opposed to food gathering, thermo-
regulation, etc.) and that are not sexually dimorphic (so are
not related to sexual selection), as Hieronymus et al. (2009)
did for centrosaurine nasal horns. However, in any case
social selection reduces to a kind of natural selection.
Moreover, these authors do not accurately distinguish
social selection and species recognition. They state (2009:
1394) that ‘species recognition traits are under selection only
in the earliest stages of courtship during mating’, following
West-Eberhard (1983); but species recognition is simply a
matter of possessing traits that allow an individual to
recognize others of its species, for many functions besides
breeding. They also state that ‘species recognition traits are
only expected to occur in closely related sympatric species,’
as opposed to being able to ‘diverge in allopatric isolated
populations,’ but in our view species recognition can begin
at the population level and can easily diverge in populations
of a single species, especially if the selective change is
anagenetic.
Contrary to West-Eberhard (1983), species recognition
does not entail ‘reproductive character displacement,’ or
necessarily any features that relate to mating, reproduction,
or competition among individuals of a species (Mayr, 1963).
Those other terms are the provenance of mate recognition,
social selection, and natural selection. She rightfully criti-
cizes earlier work that attributed to species recognition
many phenomena due to sexual selection or social selection
(such as the hypothesis that signal distinctiveness should be
reduced on islands and in isolated (allopatric) populations
(West-Eberhard, 1983: 165). That was sorted out with
further experimental work, but it does not nullify the con-
cept of species recognition or imply that it is indistinguish-
able from these other processes. This confusion aside, it is
possible to assess the predicted effects of species recognition
and to separate them from those of other hypotheses.
(iv) Species recognition – Under the explanation of species
recognition, bizarre structures would have no apparent
mechanical function and would not specifically evolve to
attract members of the opposite sex for mating (viz., Vrba,
1984; Paterson, 1993); rather, they make it easier for
9
‘Bizarre structures’ in dinosaurs
individuals to recognize others of the same (and different)
species. That is, the bizarre structures communicate to other
individuals a variety of possible associational cues, includ-
ing species identification, potential protection and social
habits and the appropriateness of potential mates. They are
positive indicators of beneficial social affiliations. There can
be a strong ontogenetic component to this process: young
neoceratopsians, pachycephalosaurs and lambeosaurs
lacked the extent of cranial ornaments of fully grown
individuals, although they had rudimentary development,
and it appears that in many cases these ornaments were
rather rapidly developed at or around the attainment of
adult size. Larger members of a species, whether male or
female, and whether or not socially dominant, thus advertise
their biological affiliation.
It is often difficult to differentiate among hypotheses of
species recognition, social selection and mate recognition,
even in living animals. All three are forms of intra-species
recognition, but less general and also different in critical
respects: it is first necessary to recognize other members of
the species, and then to recognize (in the right seasonal and
ontogenetic contexts, because mating in most species is not
year-round and does not involve all members of the popula-
tion) individuals that could serve as potential mates or
rivals. This is a different process than developing gender-
specific structures that assist in the specific attraction of
mates, or the repulsion of intraspecific competitors for
mates, which is the domain of sexual selection. Below we
propose some tests of the species recognition hypothesis that
distinguish it from the sexual selection hypothesis. In extinct
animals only hard parts generally provide evidence, and so
any evolutionary hypotheses must have an evidentiary basis
in preservable structures.
Dinosaurs and sexual dimorphism
Because sexual dimorphism has been so extensively invoked
to explain ‘bizarre structures’ in dinosaurs (e.g. Chapman
et al., 1997), we address it in detail here.
Saurischia
Sexual dimorphism has been proposed for several theropods
(mostly basal forms assigned to ‘ceratosaurs’) and ‘prosaur-
opods’ (a paraphyletic group of basal sauropodomorphs),
on the basis of an apparent difference between robust and
gracile forms (Colbert (1989, 1990) on Coelophysis; Raath
(1990) on Syntarsus). Differences have been noted in the
relative thicknesses of bone walls, and in the morphology of
trochanters. Unfortunately the statistical evidence that sup-
ports sexual dimorphism as an explanation for these differ-
ences is problematic. For example, Colbert (1990) produced
considerable evidence for ontogenetic change in proportions
in Coelophysis, but his inference of sexual dimorphism
(widely accepted by other workers) was based on only two
specimens. In Syntarsus, the difference between the ‘gracile’
and ‘robust’ morphs of the iliofemoralis trochanter is almost
non-overlapping with respect to the size of the bone (repre-
10 Journal of Zoology 283 (2011) 3–17
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K. Padian and J. R. Horner
sented by width of the femur head: Raath, 1990: Fig. 7.8).
The size-frequency distribution of femoral ‘morphs’ is also
non-overlapping with respect to the femoral head width
(Raath, 1990: Fig. 7.10). Simply put, there are no small
‘robust’ morphs. Moreover, these examples are not sexual
dimorphism in the sense established by Darwin (and John
Hunter before him); if valid sexually, they are simply slight
sexual differences, so they cannot be invoked to support
sexual selection.
An alternate possibility, that these features could be
ontogenetic, is suggested by Raath’s data. A broader tro-
chanter (and possibly thicker cortex, though the correlation
has not been statistically assessed) may have been acquired
by both males and females as they reached sexual maturity.
Sexual dimorphism has also been suggested for tyranno-
saurs (Carpenter, 1990; Larson, 1997), but Carr (1999) has
shown that many apparently dichotomous differences in the
craniofacial skeleton, such as numbers and forms of teeth,
are purely ontogenetic (as may be the case for Syntarsus), so
the ‘gracile’ forms are simply juveniles. We suspect that this
will hold for other dinosaurian species in which minor
variations in size and structure are found, rather than the
discrete structures specified by Darwin (1859, 1871) for true
sexual selection.
Other bizarre structures in theropods include cranial
crests (Dilophosaurus, Monolophosaurus, Cryolophosaurus)
and horns (Carnotaurus and incipient frontal structures in
allosaurids and tyrannosaurids); however, neither sexual
dimorphism nor ontogenetic maturity can yet been exam-
ined statistically for these features.
The argument about alleged gracile and robust dimorphic
adult forms follows, ceteris paribus, for the studies cited on
prosauropods by Galton & Upchurch (2004a: p. 257), who
provided no statistical demonstration of dimorphism, and
by Weishampel & Chapman (1990), who reached inconclu-
sive results for Plateosaurus.
Ornithischia
Sample sizes in species of stegosaurs, ankylosaurs, pachyce-
phalosaurs and most ornithopods are too small to test the
hypothesis of sexual dimorphism; it has been proposed for
hadrosaurs and ceratopsians. Goodwin (1990) noted that
the sample of pachycephalosaurs was too small to permit
statistical evaluation of sexual dimorphism, and Goodwin &
Horner (2004); Horner & Goodwin, (2009) showed that
most observed variation was ontogenetic, based on indepen-
dent analysis of stage of maturity using the degree of fusion
of the cranial sutures and the progressive growth and
reduction of specific cranial features.
Hadrosaurs
Sexual dimorphism in hadrosaurs has long been accepted by
authors (e.g. Davitashvili, 1961; Hopson, 1975; Molnar,
1977; Weishampel, 1997; Carrano, Janis & Sepkoski, 1999);
the supporting evidence can be traced almost entirely to
Dodson’s (1975) study of two genera of lambeosaurine
c 2010 The Zoological Society of London
K. Padian and J. R. Horner
hadrosaurs. Dodson’s morphometric analysis suggested
that ‘procheneosaurs’ were merely juveniles of larger spe-
cies, and he reduced three genera and 12 species to two
genera (Lambeosaurus and Corythosaurus) and three species.
In these three species he thought he could detect sexual
differences in some cranial characters, although not at all in
postcrania; and no signal was found in most cranial char-
acters. This is a problem because there is no independent
means to correlate size with age, or to identify age of a
specimen on the basis of other evidence. Evans & Reisz
(2007) have shown that this variation is ontogenetic or
characterizes chronospecies that do not overlap with each
other temporally. And moreover, these are only slight
proportional differences, not discrete structural ones.
Ceratopsians
As You & Dodson (2004) note, presumed sexual differences
have been postulated in Protoceratops andrewsi (Dodson,
1976) and (less independently) Protoceratops hellenikorhinus
for both cranial and post-cranial features, and some features
of the frill are dimorphic. However, as noted above, the
extent of variation in the supposedly dimorphic features was
statistical (as opposed to presence/absence features of true
dimorphism), and although they may have supported more
conspicuous sexually dimorphic features in soft part anat-
omy that is not preserved, the statistical argument on the
basis of hard parts is insufficient. The kind of variation
appears much more akin to the sort of differences that
characterize male and female crocodiles, which differ from
each other mainly at adult size, where it is mostly a matter of
relative robusticity (Webb et al., 1978; Chabreck & Joanen,
1979).
If dimorphism were important in small basal ceratop-
sians, it should be emphasized or at least detectable in larger,
more derived forms, but this does not seem to be the case.
Lehman (1990) suggested a pattern of sexual dimorphism in
Chasmosaurus and related species that could be traced
through later ontogeny, but the small sample sizes, incom-
plete preservation, and lack of association of much of this
material, as Lehman noted, makes it difficult to evaluate
hypotheses about sexual differences, even if they are ac-
cepted. Ryan et al.’s (2001) study of a ceratopsian bone bed,
where dimorphism could be presumed to emerge, turned up
no significant patterns. A recent review of Ceratopsia
(Dodson et al., 2004) did not accept sexual dimorphism as
a general feature in this clade of dinosaurs. Soft-part
features and behaviors that are not preserved in extinct taxa
may well have contributed to sexual selection (e.g. Sampson,
1997). However, to invoke them for extinct groups of
dinosaurs is outside the pale of homological and analogical
comparison.
As for fossil birds, which are dinosaurs, we have almost
no information about dimorphism; long tail feathers in the
basal avialian Confuciusornis are suggestive (Chiappe et al.,
1999), but this is not enough to establish evolutionary
polarity. Because dimorphism (and not just inter-sexual
difference) is generally low in other reptiles (Fig. 5), the
Journal of Zoology 283 (2011) 3–17
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c 2010 The Zoological Society of London
‘Bizarre structures’ in dinosaurs
(Sometimes, independently) (Rarely)
Non-avian dinosaurs Aves
(No) (No)
Squamata Crocodylia
Sphenodontida
(No)
(No)
Chelonia
Dimorphism in ‘bizarre’ skeletal structures?
Figure 5 We ask whether there is any support in the extant phyloge-
netic bracket of extinct dinosaurs for a broader distribution of
dimorphism in ‘bizarre’ skeletal structures. This question would not
be affirmatively answered by size dimorphism (as in crocodiles and
some other reptiles), nor by soft part structures (as in birds) or
behaviors, which cannot be assessed in fossils. The answer is
ostensibly negative.
EPB does not support sexual dimorphism in non-avian
dinosaurs on the grounds of homological comparison.
Comparative tests of the mate
competition and species recognition
hypotheses
Vrba (1984) used the example of degree of horn differentia-
tion, which is usually greater in alcelaphine bovids (harte-
beest, wildebeest, etc.) than in the related aepycerotines
(impalas), to suggest an explanation for the greater species
diversity through time of the former clade. Sampson (1999)
suggested that sexual selection, not just natural selection,
could be the motor of enhanced diversity in certain sub-
clades over others. He proposed a Mate Recognition Hy-
pothesis (MRH) by which selection for positive recognition
of mates could lead to increased differentiation of popula-
tions and eventually greater rates of speciation in some
lineages over others. This idea has a strong backing in recent
research on the value of sexual selection in promoting
differentiation of populations in a single species (e.g. An-
dersson, 1994; Price, 1998; Mendelson & Shaw, 2005). But
like any other hypothesis that involves sexual selection, a
degree of sexual dimorphism is required that is not found in
dinosaurs.
We propose that species recognition is a simpler and more
general explanation for the patterns seen in the distribution
of bizarre structures in dinosaurs. Structures that promote
species recognition allow individuals of a single species to
recognize each other and distinguish conspecifics from
members of other species. Advantages include banding
together for protection from predators, parental care and
the possible location of mates. As explained above (Display:
Intraspecific), this is a broader and more hierarchical func-
tion than that proposed by the Mate Recognition
11
‘Bizarre structures’ in dinosaurs
Figure 6 Expected differences in macroevolutionary patterns of mor-
phology between regimes governed by natural selection (or Fisherian
sexual selection) and species recognition. Although these are ex-
tremes of a continuum of patterns, it would be expected that natural
selection would cause more or less linear morphological trends
through time as the function of a structure improved. Sexual selection
might also direct the orthal elaboration of a difference into a morpho-
logical trend. But when species recognition is selected for, the object
is to be recognizably different, not necessarily functionally better or
more visually elaborate; so a range of morphology is possible, as long
as the result is divergence. From Main et al. (2005).
Hypothesis, and it does not require sexual dimorphism. It
can also involve many other kinds of cues than visual, let
alone those related to bizarre structures. The fact that these
various functions exist apart from simple mate recognition
is witnessed by the appearance of bizarre structures, often in
incipient form, in individuals not involved in mating at all.
If species recognition has been important in influencing
macroevolutionary trends, it should have some empirical
tests by which its effects can be differentiated from those of
other hypotheses. We propose two.
First, the pattern of diversification of bizarre structures in
clades should be relatively random: it should not show
trends that could ostensibly be related to selection (merely
size-related change would not qualify). An example, neces-
sarily simplified, is presented in Fig. 6. In the diagram at left,
the pattern of change documented through time shows clear
directional trends. This kind of change is readily explained
by selective forces, whether natural or sexual. The standard
model is of variation in populations, followed by directional
selection. This can represent improvement of a function
(natural selection) or continued trends in mate preference
(sexual selection; runaway sexual selection is an extreme
condition). A gradation of forms is expected both within
and among lineages: gradual improvement is expected in a
single lineage, whereas adaptive divergence (for ecological
or sexually selective reasons) should characterize differences
among lineages.
In the diagram at right in Fig. 6, however, there is no
obvious trend in evolutionary change; the only objective of
evolutionary change is to make a lineage different from
other closely related lineages (e.g. Figs 3 and 4; Main et al.,
2005: fig. 10). This pattern represents what would be more
likely expected from the species recognition model. The
direction and degree of difference are not important or
predictable; not all possible dimensions of morphospace
12 Journal of Zoology 283 (2011) 3–17
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K. Padian and J. R. Horner
are expressed. Taxonomic diversity is not necessarily higher
under either model.
Second, there could be evidence that at some point,
several closely related species with divergent bizarre struc-
tures lived at the same time in environments that at least
partly overlapped. In other words, several contemporaneous
sympatric, parapatric, or partly allopatric species existed
when these lineages were diverging. These differences might
have been positively selected as a means to reinforce
associations (including mating) with appropriate conspeci-
fics. However, lineages may also continue to diverge in
isolation from others simply because this kind of evolution-
ary change follows a natural flexibility of phenotype. So,
white-crowned sparrows diverge at the local populational
level at a very rapid rate, changing songs in ways instantly
recognizable to human birdwatchers as well as to the birds
themselves (Baptista, Bell & Trail, 1993; Bell, Trail &
Baptista, 1998). These songs both reinforce populational
identity and allow mate recognition. But the populations
may not overlap geographically to any great extent. Drift
may also play an important role, especially in small popula-
tions with some isolation (Mayr, 1963; Eldredge & Gould,
1972). Many evolutionary changes occur in lineages because
certain organisms have the evolutionary ‘habit’ of changing
regularly, not because they are adjusting to myriad contin-
uous demands of natural or sexual selection. Female pre-
ferences can change quickly, and even ‘anticipate’ desirable
variations that later appear in males (Futuyma, 2009).
In this way, we predict that the species recognition
hypothesis can account for both the differentiation of
related sympatric species and the anagenetic change in
lineages that may indeed characterize much of dinosaurian
evolution, including putative ontogenetic stages and sexual
dimorphs (e.g. Evans, 2007).
Discussion: explaining bizarre
structures in dinosaurs
Morphological diversification in the bizarre structures of
dinosaurs does not seem to show clear patterns of direc-
tional evolution within clades. To date, no satisfactory
adaptive explanation has been proposed and tested for the
evolution of bizarre structures in any dinosaurian clade (not
simply an individual species). The most recent phylogenetic
analyses of these clades do not reveal trends in the morphol-
ogy of these structures that indicate any directionality that
can be attributed to adaptive improvement or sexual selec-
tion (Weishampel et al., 2004). We stress that this does not
deny the importance of mechanical adaptation, sexual selec-
tion, or any other macroevolutionary process in dinosaurs; it
simply concludes that to date there is no evidence that it has
shaped any bizarre morphology in a clade. The fossil record
(like the living record) provides only a sample of the
diversity that has existed, and our phylogenetic reconstruc-
tions would be very different with a different or more
complete sample.
The second test of the Species Recognition model sup-
poses that several contemporaneous lineages in a clade with
c 2010 The Zoological Society of London
K. Padian and J. R. Horner ‘Bizarre structures’ in dinosaurs

Table 2 Contemporaneous occurrences of several species of related dinosaurs with ‘bizarre’ structuresa
Early Jurassic: Cryolophosaurus (Sinemurian-Pliensbachian, Ant), Dilophosaurus (Sinemurian-Pliensbachian, NAm, China)
Middle Jurassic: Stegosaurus, Lexovisaurus, ?Omosaurus (Bathonian-Callovian, Brit); Lexovisaurus (Callovian, Fr, Ger); Huayangosaurus,
unidentified stegosaur (Bathonian-Callovian, China)
Late Jurassic: Five stegosaur species (Morrison Fm., Utah), four stegosaur species (Morrison Fm., Colorado), five stegosaur species (Morrison
Fm., Wyoming) (all Kimmeridgian-Tithonian, NAm); stegosaurs, not diverse (mostly Dacentrurus armatus) (Oxfordian-Kimmeridgian, Brit;
Kimmeridgian-Tithonian, Fr, Port); at least four stegosaur species (?Oxfordian, Upper Shaximiao Fm., China)b
Early Cretaceous: Six named ankylosaur species (Cambridge Greensand, Brit), two ankylosaur species (Wessex Fm., Brit) (late Aptian).
Many species of avialians (Jiufotang and Yixian Fms., China).c
Late Cretaceous: Five neoceratopsian (‘protoceratopsids’ and relatives) species (?mid-Campanian, Djadokhta Fm., Asia).
At least four ankylosaur species, at least 10 hadrosaur species, four pachycephalosaur species, at least 10 neoceratopsian species
(Campanian, Dinosaur Park Fm., NAm);d equal or less diversity of these taxa in coeval and slightly younger formations such as the Bearpaw
Shale and the Horseshoe Canyon Fms.;e at least five pachycephalosaurs, at least three neoceratopsians (Maastrichtian, Hell Creek and Lance
Fms., Montana and neighboring states)
a
All information from Chapter 24 of Weishampel et al. (2004). Note that even though several related taxa may be present in a formation, they have
not necessarily been identified from the same localities or stratigraphic equivalences within formations. For implications, see discussion of Fig. 7
in the text.
b
The dominance of stegosaur diversity has come to light mostly in the past few decades. However, in some formations, non-‘bizarre’ dinosaurs,
such as theropods, sauropods and iguanodontids, are more diverse in the Middle and Late Jurassic.
c
Ankylosaurs replace stegosaurs in the Early Cretaceous as the most diverse ‘bizarre’ dinosaur clade. However, the non-‘bizarre’ iguanodontids
are still very diverse in places, and in China a great diversity of basal birds has been discovered in the past decade: were feathers ‘bizarre’
structures that functioned as devices of species recognition?
d
This formation also contains, among non-‘bizarre’ taxa, three tyrannosaurids, at least two ornithomimosaurs, at least three oviraptorosaurs, and
at least two dromaeosaurs.
e
The diversity of non-‘bizarre’ taxa in these formations is proportionally comparable to those in the previous note. ?Denotes uncertain identification.
Ant, Antarctica; Brit, Britain; Fm., formation; Fr, France; Ger, Germany; NAm, North America.

bizarre structures should overlap geographically to some

degree during their divergence. The geographic and tempor-
al distributions of some dinosaurian clades, including cer-
atopsians, pachycephalosaurs, lambeosaurines and
ankylosaurs, suggest that these bizarre structures appeared
in dinosaurian groups that lived at nearly the same time in
nearly the same areas (Table 2). Until recently, a salient
exception appeared to be the stegosaurs, whose low diversity
was anomalous to this general model. However, new dis-
coveries of stegosaurs have increased our knowledge of their
diversity: Carpenter (2001) estimates that at least five
stegosaur species are now known from the Morrison For-
mation of the western United States (although Galton &
Upchurch (2004b) recognize only three). This would appear
to simplify the problem, but there is an additional caveat: Figure 7 The ‘ghost’ of species recognition. Consider five species
in a lineage, distributed sequentially through time; the real extent of
the species are not all contemporaneous (K. Carpenter, pers.
temporal overlap among forms is unknown because the record
comm., 2004), and there may be geographic differentiation
is necessarily incomplete. In the figure at left, these five forms are
as well within the Morrison.
part of a single anagenetic lineage. In this case there is no evidence
The lack of contemporaneity could have several explana-
that species recognition influenced the diversification of lineages
tions, including insufficient stratigraphic sampling to estab-
(speciation), although species recognition, drift, or several other
lish that more of these species lived at the same time than it kinds of selective factors could account for the anagenetic change.
now appears. However, another approach is phylogenetic. If In the figure at right, the same five forms belong to separate line-
these five species turned out to be morphotypes of a single ages with successively ancient divergence points. This pattern
anagenetic lineage, there would indeed be no evidence for provides a basis for testing the hypothesis that species recogni-
contemporaneity. Would the hypothesis of species recogni- tion, among other factors, may have been involved in the diversifica-
tion thereby be weakened (Fig. 7, left)? In fact, such a result tion of the clade. Although as in the figure at left, no more than one of
would weaken a hypothesis of anti-hybridization, but it the five species is found at the same time as another, phylogeny
would not weaken or test the hypothesis of positive assorta- reveals their ghost lineages, and allows the inference that, for
tive mating (Paterson, 1993). However, if phylogenetic example, at time X at least four of the lineages may have been
analysis revealed that these species indeed represented contemporaneous.

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‘Bizarre structures’ in dinosaurs
different lineages, and their ‘ghost ranges’ indicated that
they must have diverged from others at an earlier time, then
at one time the test of contemporaneous species would have
been passed (Fig. 7, right). It is not impossible that such a
pattern could also indicate other processes than species
recognition, such as sexual or social selection, but in concert
with non-directional evolutionary change the indication
would be rather more strongly in favor of species recogni-
tion. Phylogenetic analysis and further biostratigraphic
sampling can test this hypothesis.
Finally, we return to the test of the Mate Recognition
Hypothesis that Sampson (1999) proposed. We found that
in every criterion, mostly related to higher rates of specia-
tion and habitat shifts, the concept of ‘species recognition’
could be substituted for the terms related to sexual selection
without any apparent difference in results. The exception
was his fourth criterion (speciation will often be correlated
with vicariance events rather than the formation of periph-
eral isolates), which we suggest is untestable in the fossil
record, and in any case would not discriminate between
sexual selection and species recognition as a cause. In
summary, the criteria for MRH and SRH are very similar
in outline; but in any given case the tests of these characters,
and perhaps the characters themselves, would be quite
different operationally for the same animals. For example,
the horns of ceratopsians might satisfy all four (five) criteria
listed above for both MRH and SRH, but would not pass
the test of high sexual dimorphism required for sexual
selection; on the other hand, they appear to pass the two
tests of the species recognition hypothesis (non-directional
variation of bizarre structures and several sympatric spe-
cies). Moreover, without a clear demonstration of sexual
dimorphism, the MRH reduces to the social selection
hypothesis (Hieronymus et al., 2009).
Conclusions
Our purpose is not to insist that species recognition has been
the only cause of the evolution of bizarre structures in
dinosaurs, nor that adaptation, social selection and sexual
selection have been unimportant in dinosaurian evolution.
We merely ask in each case: how would we test this? We
conclude that the hypotheses of mechanical function and
sexual display that have predominated for decades as gen-
eral explanations of the evolution of these structures in
dinosaurian clades are unfounded. When we test the hy-
pothesis that presumed functions of these structures have
evolved in their clades, we find no evidence; hence the
notions that these structures are ‘adaptations’ fail the
criteria proposed by evolutionary biologists (Greene, 1986;
Williams, 1992; Rose & Lauder, 1996; Padian, 2001).
Furthermore, sexual dimorphism has not been strongly
established for any bizarre structures in dinosaurian
lineages, even though mild dimorphism has been statistically
demonstrated in at least one lineage and may be plausible in
others. If criteria of sexual behavior other than those based
on sexual selection (which requires sexual dimorphism:
Darwin (1871) are to be proposed, they should be justified
14 Journal of Zoology 283 (2011) 3–17
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K. Padian and J. R. Horner
on grounds that are more stringent than weak analogies to
very different living organisms.
We stress that no evolutionary hypothesis can be re-
garded as a ‘default’ explanation (i.e. if a certain class of
explanation fails, then another one is automatically
strengthened or must be accepted by default). Hypotheses
must be independently tested, or they are not scientific. In
many or most cases, definitive tests will not be possible. We
have proposed two tests of a Species Recognition hypoth-
esis, and there may be others. In our view, most dinosaurian
bizarre structures pass these tests, but they do not pass the
tests of adaptation or of sexual display. The importance of
social selection (Hieronymus et al., 2009) remains to be
tested in dinosaurs beyond individual species. This does not
mean that these structures were not adaptive or used in
attracting mates; we simply have no evidence on these points
at present.
Our hypothesis is that the Species Recognition Hypoth-
esis is simpler and more general in explaining the evolution
of bizarre structures in dinosaurs than those of mechanical
function, social selection, or sexual selection/mate recogni-
tion. Rigorous tests of these complementary evolutionary
hypotheses should be applied to other lineages besides
dinosaurs. Bizarre structures are common in many fishes,
as well as other reptiles. In birds, sexual dimorphism,
display and selection are well-established phenomena that
have clearly had a very strong role in shaping avian evolu-
tion. The expression of bizarre structures in mammals,
notably ungulates, is entailed in a constellation of ecological
characteristics that greatly complicate their explanation
(Jarman, 1974; Perez-Barberia, Gordon & Pagel, 2002).
Finally, we emphasize that a given structure may have
several purposes, and that even in living animals it is often
difficult to determine the uses of particular structures, their
evolutionary histories, and even how the animals are com-
municating. In this respect the hypotheses of paleobiologists
are largely interpreting the shadows on the wall of Plato’s
cave. We persist in efforts to explain these structures because
they were of obvious use to their bearers, and this is in
principle discoverable.
Acknowledgments
We thank S. Bar-David, J. Brashares, V. de Buffrenil, K.
Carpenter, P. Cross, P. Dodson, J.O. Farlow, E. Hebets, T.
Hieronymus, R. Irmis, C. Janis, E. Lacey, B. Lundrigan, S.
Patek, A. de Ricqlès, M.J. Ryan, S.M. Sampson, K.M.
Scott, A.B. Shabel, L.M. Witmer and many other colleagues
and reviewers for constructive comments and suggestions,
without implying their agreement with all our points. UCB
undergraduates Jasmeet K. Dhaliwal and Sylvia Moses
provided research support. R. Irmis and A. Lee provided
technical support. This work was supported by the Univer-
sity of California Museum of Paleontology and the Com-
mittee on Research of the University of California,
Berkeley. This is UCMP Contribution No. 2012.
c 2010 The Zoological Society of London
K. Padian and J. R. Horner
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