Cladistics
Cladistics 25 (2009) 173–197
10.1111/j.1096-0031.2009.00250.x
Broken gears in the avian molecular clock: new phylogenetic analyses
support stem galliform status for Gallinuloides wyomingensis and
rallid affinities for Amitabha urbsinterdictensis
Daniel T. Ksepka
Department of Marine, Earth, and Atmospheric Sciences, North Carolina State University, Raleigh, NC 27695-8208, USA
Accepted 4 December 2008
Abstract
Galliformes (landfowl) have been the focus of numerous divergence dating studies that seek a refined understanding of the early
radiation of living birds. The Eocene fossil birds Amitabha urbsinterdictensis (Bridger Formation) and Gallinuloides wyomingensis
(Green River Formation) have been used extensively in studies dealing with the timing of evolution in crown Galliformes.
Divergence estimates from studies incorporating these fossils as calibration points suggest that multiple galliform lineages radiated
in the Cretaceous and survived the Cretaceous–Tertiary mass extinction. However, the phylogenetic position of both fossils has been
disputed, particularly with regard to crown or stem status. In order to resolve this debate, a new study of A. urbsinterdictensis and
G. wyomingensis was undertaken. Further preparation and re-examination of the A. urbsinterdictensis holotype indicates this fossil
falls outside both crown and stem Galliformes, and reveals evidence for a relationship with Rallidae (rails). In order to reassess the
status of G. wyomingensis, a matrix of 120 morphological characters was constructed by revising and expanding on previous studies.
Phylogenetic analyses using this matrix place G. wyomingensis basal to all crown Galliformes. Stem placement of G. wyomingensis is
retained and resolution is improved in combined analyses incorporating sequence data from cytochrome b, NADH dehydrogenase
subunit 2, mitochondrial control region, 12S rDNA, and nuclear ovomucoid intron G. All evidence indicates that
A. urbsinterdictensis and G. wyomingensis are inappropriate internal calibration points for Galliformes and may have contributed
to overestimation of divergence event ages. Though stem galliforms existed in the Cretaceous, the divergence of crown lineages in the
Cretaceous remains inconclusively demonstrated. Because few galliform fossils have been evaluated phylogenetically, further
investigations into the tempo of galliform evolution must await identification of proper fossil calibration points.
The Willi Hennig Society 2009.
Introduction
Molecular sequence data can be used in concert with
temporal calibration points to estimate the absolute
ages of divergences within a clade (Ayala, 1986;
Thorne et al., 1998; Sanderson, 2002, 2003; Thorne
and Kishino, 2002; Drummond et al., 2006). These
calibration points are often derived from the fossil
record, although externally derived molecular estimates
or the age of geological events resulting in isolation of
Corresponding author:
E-mail address: [email protected]
The Willi Hennig Society 2009
populations can also be used. Accurate phylogenetic
placement of fossil taxa is key to the success of
divergence-dating endeavours. One of the most important distinctions that must be made when fossils are
used as calibration points is whether a fossil belongs to
the stem group or crown group of an extant lineage.
The age of a crown group fossil (a member of the clade
uniting the most recent common ancestor of all extant
members of the clade and that ancestorÕs descendants)
provides a minimum age for divergences within that
clade. In contrast, the age of a stem group fossil (one
placed outside the crown clade) provides a minimum
age for the divergence of the clade from its sister taxon.
Historically, avian fossils have often been assigned to
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D.T. Ksepka / Cladistics 25 (2009) 173–197
extant groups without an attempt to determine their
crown or stem status. For example, compendia of
avian fossils such as Lambrecht (1933) and Brodkorb
(1964) were compiled prior to the adoption of phylogenetic systematics by ornithologists, and thus make no
distinction between stem and crown taxa. Problematic
calibrations arise either when this distinction is overlooked, or when the crown ⁄ stem status of a fossil has
not been rigorously tested (see examples in Mayr and
Mourer-Chauviré, 2003; Moyle, 2004; Parham and
Irmis, 2008).
Galliformes are part of one of the basal divergences
within extant birds. As such, they have received a great
deal of attention in molecular studies of divergence
times (Pereira et al., 2002; van Tuinen and Dyke, 2004;
Grau et al., 2005; Cox et al., 2007; Crowe et al., 2006;
Pereira and Baker, 2006). Galliformes comprises a
large radiation (281 extant species: del Hoyo et al.,
1994) including five major clades: Megapodiidae
(mound builders and allies), Cracidae (guans, chachalacas and curassows), Numididae (guineafowl), Odontophoridae (New World quail), and Phasianidae
(pheasants, partridges, grouse, turkeys and allies).
Galliformes have an abundant but patchy fossil record,
with a few species represented by multiple articulated
skeletons and many based on isolated limb bones. Two
putative crown galliform fossils have been central to
divergence-dating efforts. The first is the early Eocene
Gallinuloides wyomingensis, known from two articulated skeletons (Eastman, 1900; Dyke, 2003; Mayr and
Weidig, 2004) collected from the Green River Formation. Isolated material from other localities cannot
convincingly be assigned to Gallinuloides (see Mayr
and Weidig, 2004). The second fossil is the middle
Eocene Amitabha urbsinterdictensis, known from a
single partial skeleton collected from the Bridger
Formation (Gulas-Wroblewski and Wroblewski,
2003). Multiple molecular divergence studies have
incorporated one or both taxa as calibration points
(van Tuinen and Dyke, 2004; Crowe et al., 2006;
Pereira and Baker, 2006; Cox et al., 2007). These two
taxa are of particular importance because they are the
oldest fossils that have been used as internal calibration points for Galliformes. Gallinuloides wyomingensis
was proposed as a 55 Ma calibration point (Dyke,
2003). However, the holotype was collected from the
Fossil Butte Member of the Green River Formation
(Lance Grande, personal communication), as was the
single referred specimen (Mayr and Weidig, 2004), and
the species therefore appears to be slightly younger
than 55 Ma. Buchheim and Eugster (1998) reported an
40
Ar ⁄ 39Ar age of 50.2 ± 1.9 Ma for the Fossil Butte
Member. More recently, Smith et al. (2008) reported
a consistent but narrower 40Ar ⁄ 39Ar age of 51.66 ±
0.17 Ma for this member. Amitabha urbsinterdictensis
was collected from the Black’s Fork member of the
Bridger Formation and estimated to be 50 Ma in age
(Gulas-Wroblewski and Wroblewski, 2003). Recent
refinement of the 40Ar ⁄ 39Ar geochronology of the
Bridger Formation constrains the Black’s Fork
Member to 48.5)49.5 Ma in age (Smith et al., 2008).
The affinities of A. urbsinterdictensis and G. wyomingensis have been contentious. Various authors have placed
G. wyomingensis outside Galliformes (Eastman, 1900),
within Cracidae (Lucas, 1900; Tordoff and MacDonald,
1957; Brodkorb, 1964; Ballmann, 1969), as sister taxon to
Numididae + Phasianidae (Cracraft, 1972) or within
Phasianidae (Shufeldt, 1915). Crowe and Short (1992)
conducted a phenetic analysis that placed G. wyomingensis within crown Galliformes close to members of
Odontophoridae and Phasianidae. Dyke (2003) conducted the only previous phylogenetic analysis including
this fossil, and placed it as sister taxon to the clade uniting
Numididae, Odontophoridae and Phasianidae. Mayr
(2005, 2008) and Mayr and Weidig (2004) argued that
because G. wyomingensis lacks multiple apomorphies
shared by all extant Galliformes, the fossil instead
represents part of the stem lineage of Galliformes. These
authors provided several characters supporting this
hypothesis, but they did not present a new phylogenetic
analysis. Additionally, they suggested that erroneous
codings in the matrix of Dyke (2003) contributed to the
crown placement of G. wyomingensis. Dyke and Crowe
(2008) countered that even when information from a new
specimen (which was unavailable for Dyke, 2003
analysis) was included and corrections were made to
erroneous codings, phylogenetic analyses still placed
G. wyomingensis in the same position.
Amitabha urbsinterdictensis was originally described
by Gulas-Wroblewski and Wroblewski (2003) as a
crown galliform belonging to a clade including Odontophoridae and Phasianidae, a more exclusive placement within crown Galliformes than that proposed for
G. wyomingensis. This assignment was immediately
criticized by Mayr and Weidig (2004), who argued
that there were several flaws in the phylogenetic
analysis and that the fossil lacked convincing galliform
apomorphies. Dyke and Crowe (2008) pointed out
that in the absence of an alternative phylogenetic
hypothesis, the placement of A. urbsinterdictensis in
crown Galliformes should stand. Molecular workers
(van Tuinen and Dyke, 2004; Crowe et al., 2006;
Pereira and Baker, 2006; Cox et al., 2007) have in
general accepted a crown placement for A. urbsinterdictensis and G. wyomingensis and continued to use
these fossils as internal calibration points for Galliformes (Fig. 1).
In this paper, the relationships of A. urbsinterdictensis
and G. wyomingensis are re-evaluated in a phylogenetic
context. The character evidence for and against the
crown position of these fossils is considered alongside
new morphological characters and molecular data.
D.T. Ksepka / Cladistics 25 (2009) 173–197
175
Fig. 1. Simplified cladogram of Galliformes illustrating previously proposed fossil calibrations for divergence estimates.
Materials and methods
Osteological terminology follows Baumel and Witmer
(1993). All taxa with the exception of Paraortygoides
messelensis were coded from direct observation (see
Appendix 1). Phylogenetic analyses were conducted in
PAUP* 4.0b10 (Swofford, 2003). In all analyses, the
specified data set was subjected to a heuristic search
using 10 000 random taxon addition sequences and tree
bisection and reconnection branch swapping. Branches
of minimum length 0 were collapsed. In the combined
analysis, morphological and molecular data were
weighted equally. Specific details for each analysis are
outlined below.
Institutional abbreviations: AMNH, American
Museum of Natural History, New York, NY, USA;
MCZ, Museum of Comparative Zoology, Harvard
University, Cambridge, MA, USA; NCSM, North
Carolina Museum of Natural Sciences, Raleigh, NC,
USA; USNM, National Museum of Natural History,
Smithsonian Institution, Washington, DC, USA; WDC,
Wyoming Dinosaur Center, Thermopolis, WY, USA.
Re-evaluation of Amitabha urbsinterdictensis
Re-examination and additional preparation of the
holotype and only specimen of A. urbsinterdictensis
(AMNH 30331) was undertaken as part of the current
study. Previously unreported morphological features
and correction of misinterpretations demonstrate the
absence of any convincing galliform synapomorphies
and reveal evidence for a close relationship with either
Rallidae (rails) or a larger clade including Rallidae and
Messelornithidae (the extinct ‘‘Messel rails’’).
Gulas-Wroblewski and Wroblewski (2003, p. 1272)
listed two characters supporting referral of A. urbsinterdictensis to Galliformes: (i) ‘‘double, and open,
incisurae laterales on the sternum’’ and (ii) ‘‘incisura
capitis of proximal humerus enclosed from crus dorsale
fossae by a distinct ridge’’. The status of the first
character cannot be determined because the caudal half
of the sternum is broken off in the holotype. It appears
that a rib preserved in articulation with the sternum was
misidentified as the right trabecula lateralis, leading to
the erroneous identification of two incisurae. The second
character is present in the holotype but also occurs in
many avian groups, including Rallidae and Messelornithidae.
The phylogenetic analysis conducted by Gulas-Wroblewski and Wroblewski (2003) placed A. urbsinterdictensis as part of a clade including Odontophoridae and
Phasianidae. However, this analysis provided no test of
the higher-level relationships of the fossil because only
crown species of Galliformes were included in the
ingroup. Thus, there could be no possible outcome in
which A. urbsinterdictensis was placed with a group
other than Galliformes. The following characters reject
placement of A. urbsinterdictensis in Galliformes and
support placement close to Rallidae.
Characters supporting removal from Galliformes. 1.
Apophysis furculae weakly developed: extant Galliformes, as well as the stem fossil P. messelensis and
G. wyomingensis, possess an elongate, blade-like apophysis furculae (Fig. 2C). By contrast, the apophysis
furculae is represented only by a faintly perceptible
ridge in the furcula of A. urbsinterdictensis (Fig. 2A).
The furcula was not mentioned in the original description, although it is preserved in articulation with the
sternum.
2. Presence of a well developed processus procoracoideus: all extant and fossil Galliformes lack a processus
procoracoideus (Fig. 2F). This process is present in the
holotype of A. urbsinterdictensis (Fig. 2D), contra the
original description. A processus procoracoideus is
present in many non-galliform clades, including Rallidae (Fig. 2E) and Messelornithidae.
3. Absence of an elongate scar for accessory attachment of the tendon of m. supracoracoideus (see Baumel
and Witmer, 1993). Gulas-Wroblewski and Wroblewski
(2003) coded an elongate supracoracoideus scar as
present in A. urbsinterdictensis and extant Galliformes,
but absent in outgroups. This coding conflates the
accessory attachment scar (characteristic of Galliformes) with the tuberculum dorsale (the primary insertion
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D.T. Ksepka / Cladistics 25 (2009) 173–197
Fig. 2. Morphological features of Amitabha urbsinterdictensis compared with Rallidae and Galliformes. (A) furcula of Amitabha urbsinterdictensis;
(B) furcula of Fulica armillata (Rallidae); (C) furcula of Tetrastes bonasia (Galliformes); (D) right coracoid of Amitabha urbsinterdictensis, preserved
in cross-section; (E) right coracoid of Fulica armillata in dorsal view; (F) right coracoid of Tetrastes bonasia in dorsal view; (G) proximal right scapula
of Amitabha urbsinterdictensis preserved in articulation with coracoid; (H) proximal right scapula of Fulica armillata; (I) proximal right scapula of
Tympanuchus phasianellus (Galliformes); (J) proximal left humerus of Amitabha urbsinterdictensis in caudal view; (K) proximal left humerus of Fulica
armillata in caudal view; (L) proximal left humerus of Tympanuchus phasianellus in caudal view. Abbreviations: ap, apophysis furculae; fp, foramen
pneumaticum; pc, processus procoracoideus; sc, scar for accessory attachment of supracoracoideus; t, tubercle (see text); td, tuberculum dorsale.
site of the tendon of m. supracoracoideus, a feature
present in all extant birds). The accessory attachment
scar is absent in A. urbsinterdictensis (Fig. 2J), but
present in all extant and fossil Galliformes (Fig. 2L).
Derived characters supporting a relationship with
Rallidae. 1. Tubercle on the costal surface of the cranial
end of the scapula: this tubercle (Fig. 2G,H) was
considered an apomorphy of Rallidae by Mayr
(2006a). However, this feature is also present in Messelornithidae (Hesse, 1990; Fig. 28) and present,
although small, in some individuals of the sunbittern
Eurypyga helias (observed in AMNH 3750). The
tubercle is absent in Galliformes (Fig. 2I). As far as
can be determined, this tubercle is also absent in all
other groups not mentioned above.
2. Foramina pneumaticum absent in fossa tricipitalis
of humerus: Gulas-Wroblewski and Wroblewski (2003)
coded a large foramen pneumaticum as present in the
original description. However, the fossa tricipitalis was
still infilled with matrix when the present study was
initiated (see Gulas-Wroblewski and Wroblewski, 2003,
fig. 3A). Further preparation of the humerus reveals that
a foramen pneumaticum is absent (Fig. 2J). A foramen
pneumaticum is absent in Rallidae (Fig. 2K), but is
present in all Galliformes (Fig. 2L). The foramen
pneumaticum of the humerus is also absent in several
avian clades besides Rallidae (e.g. Palaeognathae,
Gaviiformes, Procellariiformes, Phalacrocoracidae, Recurvirostridae, Burhinidae, some Anseriformes, and the
extinct Pseudasturidae).
3. Tuberculum dorsale of humerus proximodistally
elongate and caudally projected: caudal projection and
proximodistal elongation of the tuberculum dorsale is
present in Rallidae and Messelornithidae and is also
characteristic of Psittaciformes and Columbiformes. In
contrast, the tuberculum dorsale is flush with the shaft
of the humerus and proximodistally short in Galliformes
(Fig. 2L).
Phylogenetic analysis. In order to provide a phylogenetic test of the relationships of A. urbsinterdictensis,
this taxon was coded into the matrix of Mayr and
Clarke (2003) (codings in Table 1). Phylogenetic analysis resulted in two most parsimonious trees, both
placing A. urbsinterdictensis as the sister taxon to
Rallidae. The strict consensus of these trees is shown
in Fig. 3. Relationships among the remaining taxa are
unchanged from those reported by Mayr and Clarke
(2003). A core Gruiformes clade uniting Psophiidae,
Gruidae, Rallidae, and A. urbsinterdictensis is supported by one unambiguous synapomorphy (deeply
excavated recessus caudalis fossa of pelvis), and the
clade Rallidae + A. urbsinterdictensis is supported by
one unambiguous synapomorphy (absence of foramen
pneumaticum in fossa pneumotricipitalis of humerus).
Several characters presented above as evidence against
a relationship with Galliformes (presence ⁄ absence of
processus procoracoideus) or in favour of a relationship with Rallidae (tubercle on coastal surface of
scapula) are also consistent with the topology recovered, but were not included in the matrix of Mayr and
Clarke (2003).
Although this analysis supports a relationship between A. urbsinterdictensis and core Gruiformes and
refutes a placement within Galliformes, it does not fully
resolve the phylogenetic position of the fossil, particularly whether or not it belongs to crown Rallidae. A
meaningful investigation of the fine-scale relationships
of this fossil must await further work. Relationships
D.T. Ksepka / Cladistics 25 (2009) 173–197
177
among the taxa forming the traditional order Gruiformes remain controversial, with significant conflict
between recently proposed phylogenies (compare Livezey and Zusi, 2006, 2007; Fain et al., 2007; Hackett
et al., 2008). Additionally, the extinct rail-like Messelornithidae, a clade whose relationships have been
debated (Hesse, 1990; Livezey, 1998; Mayr, 2004), needs
to be considered. In light of these issues, a full
consideration of the phylogenetic relationships of
A. urbsinterdictensis should take the form of a combined
analysis using a larger sample of taxa and attention to
the substantial fossil and subfossil record of Gruiformes
(e.g. Cracraft, 1973; Olson, 1977; Worthy and Holdaway, 2002; Livezey, 2003; Mayr, 2006a; Steadman,
2006). The focus of the present work is to test whether
or not A. urbsinterdictensis shares relationships with
Galliformes, and all evidence suggests it does not.
Phylogenetic analyses including Gallinuloides
wyomingensis
Fig. 3. Strict consensus of two most parsimonious trees (TL = 826)
from analysis including Amitabha urbsinterdictensis in the phylogenetic
matrix of Mayr and Clarke (2003). RI = 0.479, RC = 0.156.
In order to test of the relationships of G. wyomingensis, the matrix of Dyke et al. (2003) was modified and
expanded. This matrix contains the same 102 characters
Dyke (2003) used in the analysis placing G. wyomingensis within crown Galliformes, but includes a denser
sampling of ingroup taxa. Three separate analyses were
conducted to explore the effects of additional characters
and outgroup taxa added in this study, the inclusion of
molecular data, and updated codings for G. wyomingensis. Analyses were performed (i) using the original
matrix of Dyke et al. (2003) with corrections listed by
Dyke and Crowe (2008) and incorporating additional
codings from restudy of G. wyomingensis, (ii) using an
expanded morphological character set compiled for the
present study, and (iii) using the expanded morphological character set plus molecular sequence data from
five genes. Methods are outlined below.
Outgroup taxa. Broad consensus from molecular
and morphological analyses supports a sister group
relationship between Galliformes and Anseriformes,
uniting these groups in the clade Galloanserae
(Cracraft, 1988; Sibley and Ahlquist, 1990; Livezey,
1997; Groth and Barrowclough, 1999; Cracraft and
Clarke, 2001; Mayr and Clarke, 2003; Livezey and
Zusi, 2006, 2007). All recent large-scale studies of
avian phylogeny identify the split between Neognathae
and Palaeognathae as the basal divergence within
extant birds, and in turn identify Galloanserae as the
basalmost divergence within Neognathae (Groth and
Barrowclough, 1999; Mayr and Clarke, 2003; Cracraft
et al., 2004; Ericson et al., 2006; Livezey and Zusi,
2006, 2007; Hackett et al., 2008). Given this phylogenetic framework, outgroup taxa from Palaeognathae
and Anseriformes were used to polarize morphological
and molecular characters.
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D.T. Ksepka / Cladistics 25 (2009) 173–197
Table 1
Character codings for Amitabha urbsinterdictensis for the matrix of Mayr and Clarke (2003). All other characters were coded ‘‘?’’
Character
Coding
56
0
63
0
67
0
68
0
72
0
75
0
Anseriformes includes three major clades: Anhimidae
(screamers), Anseranatidae (magpie goose), and Anatidae (ducks, geese and swans). The analysis of Dyke
et al. (2003) included two species (Anhima cornuta and
Chauna torquata) representing Anhimidae and one
species (Anseranas semipalmata) representing Anseranatidae. Representatives of Anatidae were also coded in
the matrix of Dyke et al. (2003), but these taxa were
excluded from phylogenetic analysis in that study.
Therefore, only Anhimidae and Anseranatidae were
included in the reanalysis of the Dyke et al. (2003) data
set. In the new morphological analysis and combined
analysis, Anas platyrhynchos was included to represent
Anatidae. Three representatives of Palaeognathae were
newly added to the new morphological matrix. Tinamus
major and Eudromia elegans were chosen as representatives of Tinamidae. Codings from two species of
Lithornis (L. celetius and L. promiscuus) were combined
to form a single supraspecific terminal (Lithornis)
representing Lithornithidae. Lithornis was selected as
the root of the tree.
Ingroup taxa. A total of 56 ingroup species were
included. Dyke et al. (2003) coded several taxa at the
genus level, using multiple species to code a single
taxon. In the present study, a single species was
selected for each taxon to facilitate combined analysis
incorporating molecular data. Six species (Perdicula
asiatica, Odontophorus stellatus, Agelastes meleagrides,
Haematortyx sanguiniceps, Galloperdix spadicea and
Philortyx fasciatus) were omitted because sufficient
skeletal material was unavailable for study. The fossil
galliform P. messelensis, known from three specimens
from the Eocene of Messel (Germany), was added to
the matrix. Otherwise, ingroup sampling mirrors that
of Dyke et al. (2003). Specimens examined are listed in
Appendix 1.
Morphological characters. All osteological character
codings for extant taxa were re-examined by direct
observations from skeletal material. Integumentary
characters were re-examined by observations of skins
for 23 taxa and verified in the literature for all species
(Carroll, 1994; Elliott, 1994; del Hoyo, 1994; de Juana,
1994; Martı́nez, 1994; McGowan, 1994; Porter, 1994;
Jones et al., 1995; Madge and McGowan, 2002).
Codings for G. wyomingensis are based on restudy of
MCZ 342221 and the description and images of the
new specimen (WDC-CGR-012) provided by Mayr
and Weidig (2004). Codings for P. messelensis were
taken from descriptions and figures (Mayr, 2000,
2006b).
76
1
77
0
78
0
90
0
92
1
93
1
94
1
95
1
96
0
Mayr (2008) pointed out several erroneous codings in
the Dyke et al. (2003) matrix, which were corrected.
Additional problematic codings and characters were
uncovered in the present study. Thus, many characters
were modified, rescored, or excluded because they were
uninformative for Galliformes, because within-species
variability precluded reliable coding, or because codings
could not be replicated from descriptions. A total of 60
characters were added to the matrix. The expanded
morphological matrix contains 120 characters. Character definitions are presented in Appendix 2, and changes
from the original codings are marked in Appendix 3. In
order to facilitate comparisons with previous works
referencing the Dyke et al. (2003) matrix, characters
from that study are denoted with the prefix D and their
original character number. For example character 17
(D22) represents character 17 of the current study and
character 22 of Dyke et al. (2003).
Molecular data. Sequences for the mitochondrial
control region (CR), 12S rDNA (12S), cytochrome b,
and NADH dehydrogenase subunit 2 (ND2) and
nuclear ovomucoid G intron (OVO-G) were obtained
from GenBank. Sources for molecular data are cited in
Appendix 1 and largely follow those of Crowe et al.
(2006). However, in a few cases more complete
sequences were available for the current study (e.g.
Cyrtonyx montezumae and Oreortyx pictus ND2
sequences from Cox et al., 2007). Sequences were
aligned in ClustalX 1.83 (Thompson et al., 1997).
Following Crowe et al. (2006), multiple combinations
of gap opening and extension penalties were used to
align 12S and CR and the preferred alignment was
subsequently inspected and adjusted manually. Regions
of ambiguous alignment (regions where alternate gap
opening and extension penalties resulted in different
alignments) were excluded. All sequences were concatenated and added to the morphological data set for the
combined analysis. Exclusion sets correspond to positions 2864–2903, 3050–3075, 3140–3145, 3344–3352,
3625–3627, and 3761–3791 in the concatenated data set
for 12S, and positions 4006–4243, 4469–4488, and
4847–5142 in the concatenated data set for CR. The
combined data set can be accessed at TreeBASE
(http://www.treebase.org: accession no. SN420320561).
Analysis 1: Revised codings for Gallinuloides wyomingensis. In the first analysis, the effect of updating the
original codings used to place G. wyomingensis in crown
clade Galliformes was assessed. The original matrix of
Dyke et al. (2003), not including the new characters
D.T. Ksepka / Cladistics 25 (2009) 173–197
added in the current study, was utilized. Corrections for
characters 37, 86, and 101, discussed by Dyke and
Crowe (2008), were made to other taxa in the matrix,
although only presence or absence (not number) of
spurs was coded for character 86. The revised codings
for G. wyomingensis provided by Dyke and Crowe
(2008) were used, with six additional changes made
based on restudy of the material.
D1: Rostrum coded ‘‘long and shallow’’ (changed
from ‘‘?’’). The beak is complete in both specimens and
is dorsoventrally shallow (the ‘‘deep’’ condition is
present in Odontophoridae and some Cracidae).
D2: Serrations on lower jaw coded ‘‘?’’ (changed from
‘‘absent’’). These serrations are part of the rhamphotheca and thus presence or absence cannot be confirmed
in the fossil specimens, which do not preserve soft tissue.
D39: Foramen pneumaticum on dorsal surface of
coracoid coded ‘‘absent’’ (changed from ‘‘present’’). The
referred specimen of G. wyomingensis (WDC-CGR-012)
exposes the dorsal face of the coracoid, which lacks this
foramen (Mayr and Weidig, 2004).
D61: Incisura capitis of humerus coded ‘‘not enclosed
distally by a ridge’’. Dyke (2003) coded this ridge
present, and Dyke and Crowe (2008) changed the coding
to ‘‘2’’. Because no state 2 was defined for this character,
it is unclear if the intention was to retain the original
coding, change it, or denote uncertainty. Although the
proximal end of the humerus in MCZ 342221 is
damaged, the relevant area is intact and the ridge is
coded absent in the present study.
D83: Plantar side of articular surface of trochlea
metatarsi III coded ‘‘?’’ (changed from ‘‘distinctly
asymetrical’’). Neither specimen of G. wyomingensis
exposes the planter side of this trochlea.
D85: Trochleae of tarsometatarsus coded ‘‘splayed’’
(changed from ‘‘pinched together’’). See further discussion in Mayr and Weidig (2004).
All multistate characters were considered ordered,
following the methods of Dyke et al. (2003), with two
exceptions: character D92 (tail moult: irregular, centripetal or centrifugal) and character D93 (tail shape:
rounded, wedged or vaulted) were considered unordered
because the states of these two characters do not
describe a natural transition.
Phylogenetic analysis as described above resulted in
729 most parsimonious trees of 629 steps. In the strict
consensus tree (Fig. 4), G. wyomingensis is placed outside
of crown clade Galliformes as part of the galliform total
group Pangalliformes (sensu Clarke, 2004). The remainder of the tree is less resolved than, but largely congruent
with, the strict consensus tree from the original analysis of
Dyke et al. (2003). Two notable difference occur: the
clade uniting Odontophoridae, Old World quail, partridges, francolins, Galloperdix, and Ithaginis is not
recovered in the reanalysis, nor is Lophophorus recovered
as the sister taxon to Tetraoninae.
179
Fig. 4. Results of analysis 1, utilizing revised codings for the Dyke
et al. (2003) morphological matrix. Strict consensus of 729 most
parsimonious trees (TL = 629). RI = 0.631, RC = 0.112.
Analysis 2: Expanded morphological data set. The
expanded morphological analysis employed the character matrix presented in Appendix 2. Six characters were
considered ordered (see Appendix 2). The expanded
morphological analysis resulted in 1399 most parsimonious trees of 399 steps.
In the strict consensus tree (Fig. 5), G. wyomingensis
and P. messelensis are found to be sister taxa, and are
placed outside crown Galliformes.
Within crown Galliformes, four of the five major
clades supported by molecular analyses are recovered:
Megapodiidae, Cracidae, Numididae, and Odontophoridae. Odontophoridae is part of a polytomy
including many phasianid taxa, rendering Phasianidae
paraphyletic. Several previously supported subclades
of Phasianidae are recovered, including Gallininae
(junglefowl, bamboo partridges, and francolins), Pavoninae (peafowl and allies), Meleagridinae (turkeys),
and Tetraoninae (grouse). However, Coturnicinae (Old
World partridges, Old World quail, and spurfowl) and
180
D.T. Ksepka / Cladistics 25 (2009) 173–197
Fig. 5. Results of the expanded morphological analysis. Strict consensus of 1399 trees (TL = 399). Branch support values (Bremer, 1994) are
indicated below branches, and bootstrap support values are indicated above branches. RI = 0.781, RC = 0.261.
Phasianidae (pheasants) are not recovered. The strict
consensus from the morphological analysis differs
from the strict consensus tree reported by Dyke et al.
(2003) in several important aspects: (i) Numididae are
monophyletic, (ii) the two extant species of Meleagris
(wild and ocellated turkey) are monophyletic, (iii)
Polyplectron is included in Pavoninae, (iv) Gallininae
are monophyletic, (v) a francolin-spurfowl clade
(Pternistis + Francolinus) is not recovered, and (vi) a
clade uniting francolins, New World quail, Old World
quail and partridges is not recovered. In agreement
with Dyke et al. (2003), Tetraoninae (grouse) are
D.T. Ksepka / Cladistics 25 (2009) 173–197
monophyletic, but Lophophorus is not recovered as
sister taxon to Tetraoninae.
Analysis 3: Combined morphological and molecular
data set. The combined analysis resulted in 34 most
parsimonious trees of 11 788 steps.
In the strict consensus tree (Fig. 6), G. wyomingensis
and P. messelensis are recovered as sister taxa, and
181
placed outside crown Galliformes. The strict consensus
tree from the combined analysis is significantly more
resolved than the strict consensus tree from the morphological analysis. Megapodiidae, Cracidae, Numididae, Odontophoridae, and Phasianidae are all recovered
as monophyletic and are found to form successive
divergences within crown Galliformes. Within Phasiani-
Fig. 6. Results of the combined analysis. Strict consensus of 34 trees (TL = 11 788). Branch support values (Bremer, 1994) are indicated below
branches, and bootstrap support values are indicated above branches RI = 0.476, RC = 0.146.
182
D.T. Ksepka / Cladistics 25 (2009) 173–197
dae, monophyletic Arborophilinae, Coturnicinae, Gallininae, Pavoninae, and Tetraoninae (taxonomy sensu
Crowe et al., 2006) are recovered. A turkey clade was
also recovered and is labelled Meleagridinae in the
result, although it should be noted that Meleagridinae
sensu Crowe et al. (2006) also includes Perdix perdix.
Phasianinae sensu Crowe et al. (2006) was not recovered, as Ithaginis cruentus, Pucrasia macrolopha, and
Lophophorus impejanus do not form a clade with the
remaining pheasants.
Summary of results. All three analyses support placement of G. wyomingensis outside crown Galliformes. In
both the expanded morphological result and the combined result, six unambiguous synapomorphies unite all
extant Galliformes to the exclusion of G. wyomingensis:
30. Scapus claviculae of furcula narrow.
33. Spina interna of sternum present (the primitive
condition is present in P. messelensis, though this
character could not be scored for G. wyomingensis).
38. Apex carinae of sternum shifted caudally.
47. Cotyla scapularis of coracoid shallowly excavated.
58. Incisura capitis of humerus enclosed distally by a
ridge.
62. Spatium intermetacarpale of carpometacarpus
wide.
Three additional unambiguous synapomorphies unite
Cracidae and the clade Numididae + (Odontophoridae + Phasianidae) to the exclusion of G. wyomingensis, further refuting the use of this fossil as a calibration
point for the divergence between these two clades.
8. Ectethmoid highly reduced or lost.
11. Processus postorbitalis greatly elongated.
29. Furcula V-shaped.
The synapomorphies discussed above closely parallel
a list of characters provided by Mayr and Weidig (2004)
to support stem status for G. wyomingensis (seven of
those characters listed were included in the analysis).
The optimization of these characters in both the
morphological and combined analyses confirms their
relevance to galliform phylogeny.
Two character states were originally put forward
as synapomorphies of G. wyomingensis and the
clade Numididae + (Odontophoridae + Phasianidae):
(i) spatium intermetacarpale of carpometacarpus wide,
and (ii) trochleae of tarsometatarsus pinched together.
Both characters appear to have been miscoded based
on the two available specimens of the fossil taxon.
When these codings are corrected, the character of the
carpometacarpus actually supports excluding G. wyomingensis from crown Galliformes (Mayr and Weidig,
2004). In the present study, the trochleae in all
outgroup taxa and most crown Galliformes examined
were found to be similar in degree of separation. As
formulated here, only G. wyomingensis, P. messelensis,
and Megapodiidae exhibit the splayed condition. This
character thus does not support either a crown or
stem placement of the fossil taxa as it exhibits an
ambiguous optimization.
While the fossil taxa examined here are excluded from
crown Galliformes, the results suggest that reasonably
complete crown fossils should be identifiable to clades
with Galliformes via phylogenetic analysis. Most major
clades within extant Galliformes (Megapodiidae, Cracidae, Numididae, Odontophoridae, Gallininae, Pavoninae, Tetraoninae, Meleagridinae) can be recovered
from morphological data alone. One problematic area
occurs within the large Odontophoridae + Phasianidae
clade. Relationships in this sector of the tree are less well
resolved and in some areas incongruent with molecular
results, possibly owing to convergent similarities between New World quail, Old World quail and partridges. Combining morphological and molecular data
offers an avenue to overcome such difficulties.
Discussion
Given the relatively basal placement of Galliformes,
this clade is important to our understanding of the
radiation of living birds and patterns of avian survivorship across the Cretaceous–Tertiary (K-T) boundary.
Whether Galliformes diverged from Anseriformes prior
to the K-T extinction is not at issue, as the fossil
anseriform Vegavis iaai confirms this split did occur by
the Late Cretaceous (Clarke et al., 2005). However, the
question remains as to whether a single stem lineage of
Galliformes crossed the K-T boundary or multiple
crown lineages extend into the Cretaceous. Van Tuinen
and Dyke (2004), Pereira and Baker (2006), and Crowe
et al. (2006) all estimated that at least Megapodiidae
and Cracidae diverged during the Cretaceous. This
would imply that a minimum of four galliform lineages
survived the K-T extinction: at least one stem lineage
(the ancestral lineage of the P. messelensis + G. wyomingensis clade), a Megapodiidae lineage, a Cracidae
lineage, and the ancestral lineage of the clade Numididae + (Odontophoridae + Phasianidae).
However,
these previous estimates should now be reconsidered.
The incorporation of A. urbsinterdictensis and G. wyomingensis as internal calibration points is likely to have
contributed to overestimation of the age of these
divergence events. It should be noted that Pereira and
Baker (2006) also calculated Cretaceous divergences for
Megapodiidae and Cracidae in separate analyses
utilizing a 90 Ma external calibration for the Galliformes–Anseriformes split and excluding all calibration
points based on galliform fossils. However, this oft-used
external calibration has come under heavy criticism,
both because it represents a secondary calibration
(Shaul and Graur, 2002; Graur and Martin, 2004) and
because it is based on the relatively poorly constrained
mammal–bird split (Müller and Reisz, 2005). Thus,
183
D.T. Ksepka / Cladistics 25 (2009) 173–197
revised dating studies incorporating new internal fossil
calibrations are needed to address questions pertaining
to the timing of galliform evolution. It should be
stressed that while the results of the current study show
that crown galliform divergences in the Cretaceous are
not reliably supported, they do not necessarily support
placing all such divergences in the Tertiary. Other
factors, particularly assumptions about the age of the
crown radiation of birds and computational methods
used to convert branch lengths to age estimates, also
have critical impacts on divergence estimates.
Internal calibration points derived from fossils are
becoming increasingly important to divergence-dating
efforts as the limitations of externally derived calibration
points, particularly secondary calibration points, come
to light (Shaul and Graur, 2002; Graur and Martin,
2004; Ho et al., 2008). Often overlooked are the
potential pitfalls of placing fossils in a phylogenetic
context using morphological data, and subsequently
using the same fossils to calibrate the ages of nodes in a
tree derived from only molecular data. In cases where
morphological and molecular phylogenies for a group
differ significantly, it can be difficult to justify placement
of the fossil on the molecular tree (see Parham and
Irmis, 2008). Combined analysis of morphological and
molecular data including the fossil of interest is advocated here as a way to avoid this issue. Even more
important than these methodological concerns, however, is vetting of the primary phylogenetic data for the
fossils of interest. In the case of Galliformes, efforts to
understand the timing of key divergences have been
hampered both by using fossils as calibration points
based on taxonomy alone and by flawed phylogenetic
analysis. An example of the first problem is the use of
the fossil recorded as ‘‘Gallus’’ bravardi in LambrechtÕs
(1933) review of the avian fossil record to calibrate the
divergence of extant Gallus in the study of van Tuinen
and Dyke (2004). This issue was first asserted by Mayr
(2005). A reader unfamiliar with the fossil in question
might naturally assume that it represented a close
relative of extant Gallus gallus. However, this is a
dangerous assumption both because no phylogenetic
analysis has supported such a relationship, and because
of the incompleteness of the taxon (originally known
only from a partial tarsometatarsus). Indeed, MourerChauviré (1989) presented evidence from more complete
referred specimens that ‘‘Gallus’’ bravardi represents a
relative of the extant peafowl, and reclassified the species
as Pavo bravardi.
Although the potentially deleterious effects of inappropriate fossil calibrations on divergence estimates
for Galliformes are recognized here, revised divergence
estimates are not yet attainable. A full understanding
of the timing and pattern of galliform evolution will
require inclusion of the substantial fossil diversity of
this clade. As summarized in Table 2, none of the
calibration points for divergences within crown Galliformes proposed by van Tuinen and Dyke (2004) has
been confirmed by phylogenetic analysis, and several
are incorrect, as pointed out by Mayr (2005). The
Oligocene fossil Procrax brevipes from North America
was described as an early member of the Cracidae by
Tordoff and MacDonald (1957), but Olson (1985)
noted that the fossil remains incompletely prepared,
precluding a rigorous evaluation of its affinities.
Likewise, highly complete European fossils assigned
to Schaubortyx and Palaeortyx appear to represent
crown Galliformes (Mayr and Weidig, 2004; Göhlich
and Mourer-Chauviré, 2005; Mayr et al., 2006), but
their phylogenetic relationships remain untested. When
these taxa are incorporated into phylogenetic analyses,
reliable calibration points can be identified for future
investigations into the tempo of galliform evolution.
Moreover, these fossils are crucial because a complete
picture of the early galliform radiation cannot be
obtained from dates alone. Morphological data from
stem fossils of the major galliform lineages are needed
to address ecological aspects of this radiation, because
divergence estimates for stem clades do not inform us
as to when key synapomorphies of crown clades were
acquired. As a simple but important example, features
of crown Cracidae related to arboreality may have
been acquired coincident with divergence from other
Galliformes or long after this event. While molecular
data can inform us as to when stem Cracidae diverged,
fossils are necessary to reconstruct the sequence and
Table 2
Status of previously proposed fossil calibration points for Galliformes
Fossil taxon
Proposed calibration
Correct calibration
Analysis
Paraortygoides
Gallinuloides
Amitabha
Quercymegapodius
Palaeortyx
Procrax
Schaubortyx
Pavo bravardi
Galloanserae
Phasianidae–Numididae
Phasianidae–Odontophoridae
Crown Megapodiidae
Crown Odontophoridae
Crown Cracidae
Coturnix–Gallus
Crown Gallus
Galloanserae (superceded by Vegavis)
Galloanserae (superceded by Vegavis)
Inapplicable
Galloanserae (superceded by Vegavis)
Uncertain
Uncertain
Uncertain
Crown Pavo?
Yes
Yes
Yes
No
No
No
No
No
184
D.T. Ksepka / Cladistics 25 (2009) 173–197
timing of morphological transitions along the phylogenetic interval leading to crown Cracidae. The same is true
of other clades as well: fossils have much to contribute to
the study of Galliformes.
Acknowledgements
I extend thanks to Chuck Schaff and Farish Jenkins
(MCZ) for access to the holotype of Gallinuloides
wyomingensis, and to Carl Mehling and Mark Norell
(AMNH) for access and permission to further prepare
the holotype of Amitabha urbsinterdictensis. For loans of
extant comparative material, I thank Becky Desjardins
(NCSM), Joel Cracraft, Paul Sweet, George Barrowclough and Peter Capainolo (AMNH) and Chris Milensky, James Dean and Storrs Olson (USNM). Joel
Cracraft, Estelle Bourdon and an anonymous reviewer
offered constructive criticism that significantly improved
the quality of this manuscript. I also thank Kristin
Lamm and Julia Clarke for providing feedback, and
Sterling Nesbitt for preparation of the humerus and
helpful discussions. This research was supported by
NFS grant EAR 0719758.
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–
NCSM 5118
AMNH 9358, 12926
AMNH 2769, 3438;
USNM 491345
USNM 614809
Afropavo congensis
Alectoris graeca
–
–
–
–
NCSM 10634, 10635
NCSM 13942, 13943
NCSM 13053, 16698
–
NCSM 4098
–
–
AMNH 1772, USNM
347638
AMNH 11006; 12382
AMNH 961, 3781
USNM 318741
AMNH 23081
NCSM 10050, 13965
AMNH 17339
USNM 346852
AMNH 26412, USNM
492715
AMNH 3616, 4419
AMNH1313, 3439, 3589
Arborophila torqueola
Argusianus argus
Bambusicola thoracica
Callipepla californica
Callipepla squamata
Canachites canadensis
Catreus wallichi
Centrocercus
urophasianus
Chauna torquata
Chyrsolophus amhertsiae
–
–
–
Anseranas semipalmata
Anhima cornuta
Anas platyrhynchos
Ammoperdix heyi
AMNH 5366; USNM
605023
ANMH 26697; NCSM
6190,
21870, 219134
AMNH 1766, 1767 4032
NCSM 8239
AMNH 11091, 27392,
USNM 289654
Acryllium vulturinum
–
–
AMNH 2625
Aburria aburri
Alectura lathami
Skins
Skeleton
Species
AF165466
Pereira et al. (2002)
AF536742
Garcia-Moreno et al.
(2003)
AF013760
Kimball et al. (1997)
Z48772
Randi (1996)
NC007227
Slack et al. (2007)
AM236901
Crowe et al. (2006)
NC009684
Tu et al. unpublished
data
AY140735
Pereira et al. (2002)
NC005933
Harrison et al.
(2004)
AM236889
Crowe et al. (2006)
AF013761
Kimball et al. (1999)
AF028790
Kimball et al. (1999)
AB120131
Shibusawa et al.
(2004)
AF028753
Zink and Blackwell
(1998)
AF170992
Kimball et al. (1999)
AF028792
Kimball et al. (1999)
AF230177
Lucchini et al. (2001)
AY140736
Pereira et al. (2002)
AB120130
Pereira et al. (2002)
cyt b
AF222538
Dimcheff et al. (2000)
AF028773
Zink and Blackwell
(1998)
AF028758
Zink and Blackwell
(1998)
AF222548
Dimcheff et al. (2000)
DQ768254
Crowe et al. (2006)
AF222542
Dimcheff et al. (2000)
AY140738
Pereira et al. (2002)
DQ768277
Crowe et al. (2006)
–
–
NC009684
Tu et al. unpublished
data
AY140737
Pereira et al. (2002)
NC005933
Harrison et al. (2004)
AY274051
Sorenson et al. (2003)
–
AY140740
Pereira et al. (2002)
AF536745
Garcia-Moreno et al.
(2003)
DQ768253
Crowe et al. (2006)
–
ND2
(2006)
(2006)
(2006)
(2006)
AY368067
Zhan and Zhang
(2005)
DQ834478
Crowe et al. (2006)
DQ834499
Crowe et al. (2006)
AJ297158
Lucchini et al. (2001)
–
DQ834471
Crowe et al. (2006)
DQ834475
Crowe et al.
DQ834505
Crowe et al.
DQ834513
Crowe et al.
DQ834473
Crowe et al.
–
NC009684
Tu et al.
unpublished data
–
DQ834507
Crowe et al. (2006)
DQ834524
Crowe et al. (2006)
DQ834465
Crowe et al. (2006)
–
AF165430
Pereira et al. (2002)
–
CR
AF222573
Dimcheff et al. (2000)
AY140700
Pereira et al. (2002)
DQ832102
Crowe et al. (2006)
DQ834478
Crowe et al. (2006)
–
–
AF222570
Dimcheff et al. (2000)
–
–
–
NC009684
Tu et al. unpublished
data
AY140699
Pereira et al. (2002)
NC005933
Harrison et al. (2004)
AY274004
Sorenson et al. (2003)
–
–
AF165442
Pereira et al. (2002)
AF536739
Garcia-Moreno et al.
(2003)
–
12S
DQ832080
Crowe et al. (2006)
–
AF170986
Armstrong et al. (2001)
AF170980
Armstrong et al. (2001)
–
–
AF331954
Armstrong et al. (2001)
AF170978
Armstrong et al. (2001)
–
–
–
–
–
DQ832069
Crowe et al. (2006)
–
AF170991
Armstrong et al. (2001)
–
DQ832070
Crowe et al. (2006)
–
OVO-G
Taxonomy reflects current classification of Galliformes. Following Gutiérrez et al. (2000), Tetrastes bonasia is placed in the genus Tetrastes rather than Bonasia, Canachites canadensis is
placed in the genus Canachites rather than Dendragapus, and Lyrurus tetrix is placed in the genus Lyrurus rather than Tetrao. Note also that the ocellated turkey Meleagris ocellata is placed
in the genus Meleagris rather than Agriocharis, the California quail Callipepla californica is placed in the genus Callipepla rather than Lophortyx, and Pternistis afer is placed in Pternistis, not
Francolinus. These corrections account for apparent differences in taxon sampling from previous studies.
Appendix 1 Specimens examined and GenBank accession numbers for phylogenetic matrix
188
D.T. Ksepka / Cladistics 25 (2009) 173–197
AMNH 8586, 8589
AMNH 26198, NCSM
20719, USNM 500304
AMNH 11360, 14677,
27145
AMNH 8338, USNM
557976
AMNH 26842, 28230,
USNM 345096
AMNH 3748, 27953
USNM 557976
Coturnix japonica
Crax globulosa
Crossoptilon crossoptilon
Excalfactoria chinensis
Francolinus francolinus
AMNH 13873, AMNH
18359
YPM PU 16961, 23482,
23483
USNM 336535, 391983
USNM 489418
Lagopus lagopus
–
NCSM 4096
USNM 556998
NCSM 1022
AMNH 11530
–
AMNH 340, 4040; USNM –
320690, 346905
Megapodius freycinet
Meleagris gallopavo
Meleagris ocellata
Mitu mitu
–
–
–
Margaroperdix
madagarensis
Macrocephalon maleo
Lyrurus tetrix
Lophura bulweri
–
–
–
AMNH 16532, USNM
491472
AMNH 12844, USNM
500266
AMNH 12013, AMNH
27152
AMNH 523
Lithornis promiscuus
Lophophorus impejanus
Lithornis celetius
Ithaginis cruentus
AMNH 4161, USNM
430189
USNM 319188
Guttera plumifera
Gallinuloides wyomingensis MCZ 342221
Gallus gallus
AMNH
Eudromia elegans
Cyrtonyx montezumae
NCSM 16949,
16950,
16951
–
NCSM 5822
Colinus virginianus
cyt b
–
AF028796
Kornegay et al. (1993)
AF314637
Randi et al. (2001)
AF230174
Lucchini et al. (2001)
AM236881
Crowe et al. (2006)
U90640
Bloomer and Crowe
(1998)
AM236880
Crowe et al. (2006)
L08381
Kornegay et al. (1993)
–
AY141926
Pereira and Baker (2004)
AF028775
Zink and Blackwell
(1998)
NC003408
Nishibori et al.
(2001)
NCSM 20719
AY141924
Pereira and Baker
(2004)
–
AF028794
Kimball et al. (2001)
NCSM 1045, 1056 AF068192
Cox et al. (2007)
–
NC002772
Haddrath and Baker
(2001)
NCSM 11117,
NC004575
11118
Nishibori et al. (2002)
–
AF013762
Kimball et al. (1997)
–
–
NCSM 13978,
L08376
13979, 14300 (all Kornegay et al. (1993)
wild specimens)
–
AM236883
Crowe et al. (2006)
–
AF068193
Kornegay et al. (1993)
NCSM 4484, 5676 AF230170
Lucchini et al. (2001)
–
–
Skins
Skeleton
Species
Appendix 1
(Continued)
AF394631
Birks and Edwards (2002)
AF222556
Dimcheff et al. (2002)
–
AY141936
Pereira and Baker (2004)
DQ834464
Crowe et al. (2006)
DQ834485
Crowe et al. (2006)
–
AY145308
Pereira et al. (2004)
AF222564
Dimcheff et al. (2000)
AF394621
Birks and Edwards (2002)
–
DQ834528
Crowe et al. (2006)
–
DQ834486
Crowe et al. (2006)
AJ300146
Randi et al. (2001)
DQ834479
Crowe et al. (2006)
–
DQ834487
Crowe et al. (2006)
DQ834482
Crowe et al. (2006)
–
DQ768258
Crowe et al. (2006)
AF222552
Dimcheff et al. (2000)
–
DQ834514
Crowe et al. (2006)
–
DQ834510
Crowe et al. (2006)
–
DQ834500
Crowe et al. (2006)
DQ834467
Crowe et al. (2006)
–
–
–
DQ768259
Crowe et al. (2006)
–
NC003408
Nishibori et al. (2001)
Af22576
Dimcheff et al. (2000)
12S
–
–
AF170981
Armstrong et al. (2001)
AF170976
Armstrong et al. (2001)
–
–
AY952773
Cox et al. (2007)
AY952772
Cox et al. (2007)
OVO-G
–
AF170984
Armstrong et al. (2001)
–
–
U83741
Mindell et al. (1997)
–
–
–
–
–
–
DQ832075
Crowe et al. (2006)
–
–
DQ832076
Crowe et al. (2006)
–
–
–
AF222593
Dimcheff et al. (2000)
–
–
DQ832098
Crowe et al. (2006)
–
AF222583
Dimcheff et al. (2000)
–
–
–
–
NC001323
AF170979
Desjardins and Morais Armstrong et al. (2001)
(1990)
–
–
AY952764
Cox et al. (2007)
NC002772
Haddrath and Baker
(2001)
AB073301
Nishibori et al. (2002)
–
–
AY145316
–
Pereira et al. (2002)
–
DQ834469
Crowe et al. (2006)
CR
–
–
AB086102
Wada et al. (2004)
AY141934
Pereira and Baker
(2004)
DQ768256
Crowe et al. (2006)
AY952748
Cox et al. (2007)
NC002772
Haddrath and Baker
(2001)
NC004575
Nishibori et al. (2002)
–
NC003408
Nishibori et al. (2001)
AF222545
Dimcheff et al. (2000)
ND2
D.T. Ksepka / Cladistics 25 (2009) 173–197
189
NCSM 6385
–
AMNH 16428, 16429;
USNM 289116
AMNH 1368, USNM
613959
AMNH 27055, USNM
623356
AMNH 20970
AMNH 21989, 22689,
22690
AMNH 4192, 5051
AMNH 4142, 4184, 4300
AMNH 27147
AMNH 4029
AMNH 2741
AMNH 11062, NCSM
20205
USNM 290303
AMNH 25536; USNM
430182
AMNH 5099, 26515
AMNH 5283, USNM
621934,
621935
AMNH 614, 1318, 26498
Pavo cristatus
Penelope purpurascens
Polyplectron inopinatum
Pternistis squamatus
Pucrasia macrolopha
Rheinardia ocellata
Rhizothera longirostris
Rollulus roulroul
Tetraogallus himalayensis
Tetrastes bonasia
Tinamus major
Tympanuchus phasianellus AMNH 23620, USNM
559926
Tragopan satyra
Syrmaticus reevesi
Pternistis afer
Phasianus colchicus
Perdix perdix
Ortalis vetula
NCSM 12616, AF534555
16966
Bush and Strobeck (2003)
NCSM 19106, AF068191
19107
Kimball et al. (1999)
AY637103
Pereira et al. (2002)
NCSM 10193 AF028791
Kimball et al. (1999)
NCSM 1858, AY368060
3015, 7134
Zhan and Zhang,
unpublished data
–
AF330064
Kimball et al. (2001)
–
U90635
Bloomer and Crowe (1998)
–
U90636
Bloomer and Crowe (1998)
–
AF028800
Kimball et al. (1999)
–
AF330060
Kimball et al. (1999)
–
–
NCSM 20205, AM236888
20626
Crowe et al. (2006)
NCSM 13917 AF368059
Zhan and Zhang,
unpublished data
–
AY678108
Qiang et al. unpublished
data
–
AF230165
Lucchini et al. (2001)
–
NC002781
Haddrath and Baker (2001)
L08379
Kornegay et al. (1993)
AMNH 5124, 5291, 23328; NCSM 13928
NCSM 17388
AMNH 23814
NCSM 4102,
4104
AMNH 1405, USNM
NCSM 22759
288723
Numida meleagris
Oreortyx pictus
AF165470
Pereira et al. (2002)
L08383
Kornegay et al. (1993)
AF252860
Armstrong et al. (2001)
L08384
Kornegay et al. (1993)
–
AMNH 6042, 6043
Nothocrax urumutum
cyt b
Skins
Skeleton
Species
Appendix 1
(Continued)
(2006)
(2006)
(2006)
(2006)
AF222560
Dimcheff et al. (2002)
AF222539
Dimcheff et al. (2000)
NC002781
Haddrath and Baker
(2001)
–
–
DQ768271
Crowe et al. (2006)
–
–
DQ768267
Crowe et al.
DQ768281
Crowe et al.
DQ768286
Crowe et al.
DQ768269
Crowe et al.
–
AY140749
Pereira and Baker (2004)
NC006382
Nishibori et al. (2004)
AY952749
Cox et al. (2007)
AF394614
Birks and Edwards
(2002)
AF394612
Birks and Edwards
(2002)
AY367097
Pereira et al. (2002)
AF222560
Dimcheff et al. (2000)
AF222561
Dimcheff et al. (2000)
ND2
DQ834489
Crowe et al. (2006)
DQ834483
Crowe et al. (2006)
DQ834477
Crowe et al. (2006)
–
DQ834520
Crowe et al. (2006)
DQ834492
Crowe et al. (2006)
AJ29528
Kimball et al. (2001)
DQ834533
Crowe et al. (2006)
DQ834531
Crowe et al. (2006)
DQ834490 Crowe et al.
(2006)
DQ834506
Crowe et al. (2006)
–
–
AY145312
Pereira et al. (2002)
DQ834484
Pereira et al. (2002)
DQ834495
Pereira et al. (2002)
DQ834508
Crowe et al. (2006)
AF165440
Pereira et al. (2002)
DQ834466
Crowe et al. (2006)
DQ834468
Crowe et al. (2006)
–
CR
–
–
–
–
–
–
–
AF222598
AF170985
Dimcheff et al. (2000) Armstrong et al. (2001)
AF230147
Lucchini et al. (2001)
NC002781
Haddrath and
Baker (2001)
–
–
–
–
–
–
DQ832111
Crowe et al. (2006)
DQ832109
Crowe et al. (2006)
–
AF331958
Kimball et al. (2001)
DQ832092
Crowe et al. (2006)
DQ832088
Crowe et al. (2006)
AF170983
Armstrong et al. (2001)
–
AF170982
Armstrong et al. (2001)
AY952774
Cox et al. (2007)
AF222590
Dimcheff et al. (2000)
U83742
Mindell et al. (1997)
–
–
AF170975
Armstrong et al. (2001)
AF170977
Armstrong et al. (2001)
AF170874 Armstrong
et al.
(2001)
AF170990
Armstrong et al. (2001)
–
OVO-G
–
AF165446
Pereira et al. (2002)
AF222587
Dimcheff et al. (2000)
AY952765
Cox et al. (2007)
U88017
Cooper and Penny
(1997)
AY722396
Saini et al. (2007)
12S
190
D.T. Ksepka / Cladistics 25 (2009) 173–197
D.T. Ksepka / Cladistics 25 (2009) 173–197
Appendix 2 Morphological character descriptions
Characters 35, 36, 37, 55, 67, and 109 are treated as ordered.
Characters from the Dyke et al. (2003) matrix study are denoted with
the prefix ‘‘D’’ and their original character number.
Osteology
1. (D1). Rostrum: dorsoventrally deep (0); dorsoventrally shallow (1).
After Holman (1964).
2. Beak, spatulate shape in dorsal view: absent (0); present (1).
3. (D9). Width of pila supranasalis between external nares: wide (0);
narrow (1).
4. (D10). Premaxilla, processus nasalis: divides rostral portion of
frontal (0); does not divide rostral portion of frontal (1).
5. Nasal septum: absent (0); present (1).
6. Lacrimal, processus supraorbitalis: no posterior projection into orbit,
or weak and blunt projection (0); forms a sharp spine projecting into
orbit (1). Characters 6 and 7 were split from character D16.
7. Lacrimal, facies articularis frontonasalis in dorsal view: contact with
frontal forms a straight suture (0); lacrimal occupies a notch in lateral
margin of frontal (1).
8. (D8). Ectethmoid: present (0); highly reduced or lost (1). Cracraft
(1968) provides a thorough discussion of this character.
9. (D17). Maxillopalatine shelf: absent (0); present (1). In the original
use of this character, some Megapodiidae and Cracidae were coded as
having a maxillopalatine shelf. Because the maxillopalatines approach
one another without contact and fusion in these taxa, they are coded 0
in the current study.
10. Palatine and pterygoid: fused (0); separate (1).
11. Processus postorbitalis: short (0); greatly elongated (1). This
character and characters 12–13 replace character D19. In some
representatives of Megapodiidae, the processus postorbitalis and
processus zygomaticus fail to fuse distally due to the short length of
the processus postorbitalis. In representatives of Cracidae, lack of
fusion of these processes is instead due to the shorter ossified portion
of the aponeurosis zygomatica. Scoring the length of the processus
postorbitalis and processus zygomaticus individually (this character
and character 12) is more representative of morphological variation
than coding only presence or absence of fusion.
12. Processus zygomaticus: well developed (0); absent or poorlydeveloped (1); processus zygomaticus short, but continuous with well
ossified aponeurosis zygomatica which extends anterior to near or
beyond the level of the postorbital process (2). Zusi and Livezey (2000)
evaluated the homology of cranial structures of Galloanserae and
other birds. These authors differentiated between taxa with a prominent processus zygomaticus (e.g. Tinamidae) and those with a poorlydeveloped processus zygomaticus that superficially appears elongate
due to the attached ossified aponeurosis zygomatica (e.g. many
Galliformes). These conditions are non-homologous and therefore
treated as separate character states.
13. (D19). Processus postorbitalis and processus zygomaticus (including
aponeuroses): unfused (0); fused distally (1). The character state
‘‘partially fused’’ was eliminated as it was not observed in any taxa.
14. Rounded flange projecting ventrally from dorsal margin of tympanic
region: absent or weak (0); strongly developed (1). See Fig. 7.
15. Processus basipterygoideus: long and arising posteriorly (0); short
and arising anteriorly on parasphenoid rostrum (1).
16. Quadratojugal–quadrate articulation: quadratojugal articulates at
the level of the ventral extent of the condylus caudalis (0); quadratojugal articulates well dorsal to the level of the condylus caudalis. See
Fig. 7.
17. (D22). Quadrate, processus orbitalis: short (0); long (1).
18. Mandible, processus coronoideus: absent or poorly developed (0);
strongly projected (1). After Livezey (1997), character 18.
19. Mandible, deep groove on ventral surface of symphysis: absent (0);
present (1). This character was discussed by Olson and Feduccia
(1980).
20. Mandible, fenestra mandibularis caudalis: absent (0); present (1).
191
A
B
C
D
Fig. 7. Lateral view of the skull of (A) Alectura lathami (Megapodiidae: USNM 614809); (B) Penelope purpurascens (Cracidae: USNM
613959); (C) Cyrtonyx montezumae (Odontophoridae: USNM 557976);
(D) Pucrasia macrolopha (Phasianidae: AMNH 27147) illustrating
character states for characters 1, 5, 8, 11, 12, 13, 14, and 16. Not to
scale. Abbreviations: ec, ectethmoid; fl, flange projecting from dorsal
margin of tympanic region; fp, fusion of processus postorbitalis and
processus zygomaticus; ns, nasal septum; pp, processus postorbitalis;
pz, processus zygomaticus; qj, quadratojugal-quadrate articulation.
21. Mandible, two strong grooves on ventral surface of symphysis: absent
(0); present (1).
22. Axis, foramina transversaria: absent (0); present (1).
23. Cervical vertebrae 3 and 4, bony bridge from processus transversus to
processus articularis caudalis: absent (0); present (1). Mayr and Clarke
(2003: character 52) included this character but referred only to
cervical vertebra 3. The derived state in the present conformation also
refers to cervical vertebra 4.
24. (D23). Thoracic vertebrae, lateral pneumatic fossa: present (0);
absent (1)
25. Notarium, degree of fusion of thoracic vertebrae: complete (0);
partial (1).
26. (D26). Notarium, number of incorporated vertebrae: four or less (0);
five (1).
27. Synsacrum, processes transversus of sacral vertebrae at level of
acetabulum forming dorsoventrally tall lamina that contacts the medial
margin of acetabulum: absent (0); present (1).
28. Processus uncinatus: fused to ribs (0); not fused to ribs (1); absent
(2).
29. (D27). Furcula: U-shaped (0); V-shaped (1).
30. Furcula, scapus claviculae: stout (0); slender (1). This character was
discussed by Mayr and Weidig (2004).
31. (D28). Furcula, scapus claviculae: widening towards extremitas
omalis (0); of uniform thickness (1).
32. (D29). Furcula, apophysis furculae: small or obsolete (0); pronounced projection (1).
33. Sternum, spina interna: absent (0); present (1).
34. Sternum, spina externa: absent (0); present (1).
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D.T. Ksepka / Cladistics 25 (2009) 173–197
35. (D51). Sternum, processus craniolateralis: perpendicular to carina
(0); oriented at angle of 45 degrees with respect to carina (1); parallel to
carina (2). Ordered. After Holman (1964).
36. (D52). Sternum, processus craniolateralis: short (0); moderate
length (1); long (2). Ordered. This character originally contained 4
states (Dyke et al., 2003), but three states are considered sufficient to
encompass variation in the present study. After Holman (1964).
37. (D53). Sternum, processus craniolateralis: wide (0); moderate width
(1); narrow (2). Ordered. After Holman (1964).
38. Sternum, apex carinae: extends far cranially (0); shifted caudally
(1). As discussed by Stegmann (1964) and Mayr (2006b) the derived
morphology is associated with development of an enlarged crop. See
Fig. 8.
39. Sternum, marked sulcus along cranial face of carina: absent (0);
present (1).
40. (D48). Sternum, caudal incisurae: single (0); double (1).
41. Sternum, incisurae medialis et lateralis: shallow (0); deep (1).
42. Sternum, caudal margin: wide (0); tapering (1).
43. (D32). Scapula, acromion: hooked (0); flat (1). After Holman
(1964). See Fig. 9.
44. (D34). Scapula, facies articularis humeralis: parallel to corpus
scapulae (0); acute with respect to corpus (1).
45. (D35). Scapula, dorsal surface of facies articularis humeralis:
excavated by fossa (0); fossa absent (1). After Holman (1964).
46. (D36). Scapula, fossa between acromion and facies articularis
humeralis: present (0); absent (1). After Holman (1964). See Fig. 9.
A
pc
se
ac
B
pc
si
se
ac
pc
C
si
se
ac
Fig. 8. Lateral view of the sternum of (A) Anas platyrhynchos
(Anatidae: NCSM 21934); (B) Crax globulosa (Cracidae: AMNH
20719); (C) Gallus gallus (Phasianidae: NCSM 4105) illustrating
character states for characters 33, 34, 35, 36, 37, 38, 40, and 41. Not
to scale. Abbreviations: ac, apex carinae; pc, processus craniolateralis;
se, spina externa; si, spina interna.
A
B
Fig. 9. Cranial view of the right scapula of (A) Centrocercus urophasianus (Phasianidae: USNM 492715); (B) Alectura lathami (Megapodiidae: USNM 614809) illustrating characters 43 and 46.
Abbreviations: ac, acromion; fah, facies articularis humeralis; pf,
pneumatic foramen.
47. (D37). Coracoid, cotyla scapularis: cup-like, excavated (0); shallow,
not excavated (1). Mayr and Weidig (2004) discussed this feature in the
context of galliform phylogeny.
48. (D40). Coracoid, distinct processus procoracoideus: present and
projected (0); absent (1).
49. Coracoid, foramen nervi supracoracoidei: present (0); absent (1).
50. Coracoid, blunt ventral projection at omal end, adjacent to facies
articularis clavicularis: absent (0); present (1).
51. (D39). Coracoid, distinct fossa pneumaticum on dorsal surface:
present (0); absent (1). After Holman (1964).
52. (D45). Coracoid, facies articularis sternalis: grades smoothly into
dorsal surface of shaft (0); bordered dorsally by a strong raised lip (1).
This character has been modified to reference the lip, and codings
differ markedly from the original codings of Dyke et al. (2003) for
character D45.
53. Coracoid, processus lateralis: rounded, with weak projection (0);
pointed, with strong projection (1). The shape of the processus lateralis
was divided into three characters by Dyke et al. (2003): 43, 44, 46). A
single character is considered sufficient to represent variation between
taxa here.
54. Humerus, crista bicipitalis in anterior view: rounded (0); squared (1).
This feature was discussed by Holman (1964) as well as Crowe and
Short (1992).
55. (D56). Humerus, secondary fossa pneumotricipitalis on caudal face:
rudimentary or absent (0); moderately developed (1); strongly developed, forming deep excavation (2). Ordered. This character originally
contained four states, but three are considered sufficient to encompass
variation in the present study. After Holman (1964). See Fig. 10.
56. (D60). Humerus, caudal surface, foramen pneumaticum: small (0);
large (1). After Holman (1964).
57. (D58). Humerus, elongate raised crest on shaft, distal to tuberculum
dorsale (this crest represents an accessory insertion of the tendon of m.
supracoracoideus): absent (0); present (1). See Figs 2 and 10.
58. (D61). Humerus, incisura capitis: open groove (0); enclosed distally
by a ridge (1). The derived state of this character was recognized as a
synapomorphy of crown Galliformes by Mayr (2000). See Fig. 10.
59. (D59). Humerus, distal extent of condylus ventralis in cranial view:
not markedly extended distally (0); markedly protrudes distally (1).
60. (D62). Ulna: shorter or equal to humerus in length (0); longer than
humerus (1).
61. Carpometacarpus, ventral face, proximal margin of rim of trochlea
carpalis: smoothly rounded (0); with small notch (1).
62. (D65). Carpometacarpus, spatium intermetacarpale: narrow (0);
wide (1). This character was discussed by Mayr (2000).
63. (D66). Carpometacarpus, processus intermetacarpalis: absent (0);
present and overlapping metacarpal III (1). After Holman (1964).
64. Carpometacarpus, large bony spur projecting from anterior face of
carpometacarpus: absent (0); present (1).
193
D.T. Ksepka / Cladistics 25 (2009) 173–197
A
B
C
D
Fig. 10. Caudal view of the right humerus of (A) Chauna torquata (Anhimidae: AMNH 3616); (B) Penelope purpurascens (Cracidae: USNM
613959); (C) Pucrasia macrolopha (Phasianidae: AMNH 27147); (D) Callipepla squamata (Odontophoridae: NCSM 10050) illustrating characters 55,
57, and 58. Not to scale. Abbreviations: r, ridge enclosing incisura capitis; sc, scar for accessory attachment of m. supracoracoideus ridge; sf,
secondary fossa tricipitalis.
65. Carpometacarpus, anterior face: flat or rounded (0); sharp ridge
present anteriorly (1).
66. Carpometacarpus, metacarpal III strongly twisted: absent (0);
present (1).
67. Alular digit, rudimentary claw: absent (0); very small and buttonshaped (1); claw-like (2). Ordered. This small claw is often lost during
maceration of specimens. Stephan (1992) confirmed the true absence of
this claw in several taxa. Observations from that study were used to
supplement codings from specimens preserving the claw. Specimens
that lacked a claw were coded ‘‘?’’ unless true absence could be
confirmed from Stephan (1992).
68. (D69). Pelvis, cranial margin flared lateral to the margo dorsalis:
present (0); absent (1).
69. (D71). Pelvis, canalis iliosynsacralis opens posteriorly at two large,
depressed foramina located between the iliac anterior crests and the
crista spinosa synsacra: present (0); absent (1). See Fig. 11.
70. Pelvis, anteriorly directed tab-like process placed dorsal to the
antitrochanter: absent (0); present (1). See Fig. 11.
71. Pelvis, ilia and crista spinosa synsacri: remain separate (0); fused at
dorsal margin (1).
72. (D68). Pelvis, tuberculum preacetabulare (pectineal process): long
and projected (0); small point (1). After Holman (1964).
73. Pelvis, spina dorsolateralis ilii projects as sharp mediolaterally
narrow process, adjacent to lateral margin of synsacrum and proximal
caudal vertebrae: absent (0); present (1). See Fig. 11.
74. Foramen ilioischiadicum: open caudally (0); closed (1).
75. Pelvis, recessus caudalis fossa: shallow (0); deep (1); absent (2).
76. (D74). Depth of ischium relative to the width of the synsacrum: deep
(0); shallow and wide (1).
77. Spatium ischiopubicum: dorsoventrally wide (0); dorsoventrally
narrow and slit-like (1). After Livezey and Zusi, 2006, character 1787.
78. Femur, length: shorter or equal to humerus (0); longer than
humerus (1).
79. (D77). Femur, fossa poplitea: deeply recessed with pneumatic
foramen ⁄ fossa (0); not deeply recessed, foramen variably present (1).
80. Tibiotarsus, crista cnemialis cranialis, proximal apex: flat or
rounded in anterior view (0); pointed (1). See Fig. 12.
81. Tarsometatarsus, bony canal enclosing tendon of m. flexor digitorum
longus: absent (0); present (1).
82. (D86). Tarsometatarsus, spurs in males: absent (0); present (1). This
character cannot be scored in fossil taxa given the lack of sex data for
such taxa. After Holman (1964).
83. Tarsometatarsus, foramen at distal end of shaft between trochlea
metatarsi II and III: absent (0); present (1).
84. (D82). Tarsometatarsus, relative length of trochleae: trochlea
metatarsi II and IV of similar length (0); trochlea metatarsi II
distinctly shorter than trochlea metatarsi IV. After Holman (1964).
A
B
C
Fig. 11. Dorsal view of the pelvis of (A) Alectura lathami (Megapodiidae: USNM 614809); (B) Penelope purpurascens (Cracidae: USNM
613959) and Francolinus francolinus (Phasianidae: USNM 557976)
illustrating characters 69, 70, 71, 72, and 73. Not to scale. Abbreviations: c, open contact between ilium and crista spinosa synsacri; ci,
posterior opening of canalis iliosynsacralis; f, fusion of ilium and crista
spinosa synsacri; sdi, spina dorsolateralis ilii; t, tab-like process dorsal
to antitrochanter; tp, tuberculum preacetabulare.
194
D.T. Ksepka / Cladistics 25 (2009) 173–197
A
B
Fig. 12. Anterior view of the left tibiotarsus of (A) Penelope purpurascens (Cracidae: USNM 613959); (B) Meleagris gallopavo (Phasianidae: NCSM 22760) illustrating relative proximal projection of the
crista cnemialis cranialis (character 79). Not to scale.
85. (D83). Tarsometatarsus, plantar side of articular surface of trochlea
metatarsi III: symmetrical (0); distinctly asymmetrical with lateral
ridge protruding farther proximally than medial ridge (1). General
asymmetry of trochlea metatarsi III is present in many avian clades
and so is not considered diagnostic of Galliformes. The more detailed
definition of the derived state here follows Mayr (2000), and can only
be scored in plantar view.
86. (D85). Tarsometatarsus, trochleae: splayed (0); close together (1).
Character state (1) was originally proposed as a synapomorphy uniting
Cracidae and other crown Galliformes to the exclusion of Megapodiidae. However, outgroup taxa also show state (1), resulting in a
homoplastic distribution.
87. (D87). Tarsometatarsus, length of toes relative to tarsometatarsus:
short (0); long, digit III subequal or longer than tarsometatarsus (1).
The derived state is quantified here. The coding for Lithornis is based
on measurements from Houde (1988).
88. Hallux: significantly shorter than remaining pedal digits (0); greatly
elongated, approaches or exceeds other digits in length (1).
89. (D89). Hallux: incumbent (at same level as remaining pedal digits)
(0); elevated, more proximally located than remaining digits (1).
Arthrology
90. Ligamentum postorbito-mandibulare: absent (0); present (1).
Codings follow Zusi and Livezey (2000).
Plumage
91. (D101). Integument of head: largely feathered (0); largely naked (1).
Three states were defined in the original formulation of this character,
but as no taxa were scored state 2 (totally naked) that state was
eliminated. Lophura bulweri is coded variable, as males have a largely
naked head while females have a largely feathered head.
92. Single elongate ornamental plume on head: absent (0); present (1).
93. Tuft of ornamental plumes with expanded distal ends on head: absent
(0); present (1).
94. (D98). Orbit: feathered (0); patch of bare skin surrounds orbit (1).
The definition of this character is slightly modified to reference the skin
surrounding the orbit rather than the eye-ring. The skin surrounding
the orbit of Anseranas semipalmata is surrounded by feathers posteriorly, but bare anteriorly. This taxon is coded (0) in the present study.
95. (D95). Body plumage black, spotted with white vermiculations:
absent (0); present (1).
96. Body plumage, black and white vertical barred plumage on flank:
absent (0); present (1).
97. Contour feathers, downy barbules at base: lack detachable nodes (0)
possess detachable nodes (1). Codings follow Brom (1991).
98. (D94). Wing: longer than tail (0); shorter than tail (1). Gallus gallus
is coded 0 ⁄ 1 to reflect the variation in tail length between males and
females.
99. Wing feathers: diastataxic (0); eutaxic (1). Codings follow Jones
et al. (1995) and Bostwick and Brady (2002).
100. Outermost primaries: unmodified (0); tip bowed and stiffened for
acoustic use (1). Codings follow the descriptions of del Hoyo (1994).
101. (D91). Number of tail feathers: less than 16 (0); equal to or greater
than 16 (1).
102. (D93). Tail shape: round (0); wedged or graduated (1), vaulted (2).
103. (D92). Tail feather moult: irregular or bi-directional (0); centrifugal (1); centripetal (2).
104. (D90). Tarsus: unfeathered (0); at least partially feathered (1).
105. (D97). Sexual dimorphism in plumage: absent (0); slight (1),
marked (2).
106. Integument of hatchling: downy (0); true feathers (1). The first
generation of true feathering is fully grown in megapode hatchlings
upon emergence from the nest (Clark, 1964).
Miscellaneous soft tissue
107. Fleshy, brightly coloured comb dorsal to eye: absent (0); present (1).
108. (D2). Lower beak, serrations on cutting edge of rhamphotheca:
absent (0); present (1). This character description has been modified to
refer to the rhamphotheca, because the rhamphotheca alone (not the
underlying bony mandible) is serrated in Odontophoridae
109. Filtering lamillae: absent (0); rudimentary (1); well developed (2).
Ordered. After Livezey, 1997, character 103.
110. Frontal caruncle: absent (0); present (1). This structure, also
referred to as a snood, is present in turkeys.
111. Single wattle formed by skin of neck: absent (0); present (1).
112. Paired wattles formed by skin on side of face (at least in male):
absent (0); present (1).
113. Inflatable cervical air sacs (used for display): absent (0); present (1).
114. Tracheal elongation in males: absent (0); present (1). Scorings
follow Fitch (1999). Penelope purpurascens differs from most other
members of the genus Penelope in lacking tracheal elongation and the
grouse Tetrao urogallus (not included in the present analysis) also
possesses tracheal elongations (Fitch, 1999).
115. Intromittant organ: absent (0); present (1). Codings follow Brom
and Dekker (1992) and Oliveira et al. (2004).
116. Uropygial gland: naked (0); tufted (1). Codings follow Johnston
(1988).
Eggs and reproductive behavior
117. Eggshell, pinkish brown powdery covering: absent (0); present (1).
Codings follow Brom and Dekker (1992).
118. (D99). Average clutch size: four or more eggs (0); two or three
eggs (1). For some galliform species, clutch size remains poorly
known. For example, only one nest has ever been recorded in wild
Lophura bulweri (McGowan, 1994). Although this nest contained a
single egg, this taxon is scored ‘‘?’’ since the sample size is too small
to rule out the possibility this number was the result of a single
abnormal clutch or an instance of predated eggs. In this respect,
codings are more conservative (and less complete) than those of Dyke
et al. (2003).
119. (D100). Mating system: monogamous (0); polygynous (1).
120. (D102). Incubation system: egg incubated by parents (0); egg
incubated by external means, e.g. geothermal heat or decaying
vegetation (1).
D.T. Ksepka / Cladistics 25 (2009) 173–197
Appendix 3 Morphological data matrix
Codings that differ from Dyke et al. (2003) for characters that directly overlap with that study are in bold. Symbols A = 0 ⁄ 1, B = 1 ⁄ 2.
195
196
Appendix 3
(Continued)
D.T. Ksepka / Cladistics 25 (2009) 173–197
D.T. Ksepka / Cladistics 25 (2009) 173–197
Appendix 3
(Continued)
197