Group Title: Bulletin of the Florida State Museum
Title: Osteology of living and fossil New World quails (Aves, Galliformes)
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Full Citation
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Permanent Link: http://ufdc.ufl.edu/UF00001505/00001
 Material Information
Title: Osteology of living and fossil New World quails (Aves, Galliformes)
Alternate Title: Bulletin of the Florida State Museum ; volume 6, number 2
Physical Description: 132-233 p. : tables. ;
Language: English
Creator: Holman, J. Alan, 1931-
Publisher: University of Florida
Place of Publication: Gainesville, Fla.
Publication Date: 1961
 Subjects
Subject: Quails   ( lcsh )
Skeleton   ( lcsh )
Birds -- Anatomy   ( lcsh )
Genre: bibliography   ( marcgt )
government publication (state, provincial, terriorial, dependent)   ( marcgt )
non-fiction   ( marcgt )
 Notes
Thesis: "The present paper represents in part a doctoral dissertation accepted at the University of Florida in 1961."
Bibliography: "Literature cited": p. 230-233.
General Note: Cover title.
 Record Information
Bibliographic ID: UF00001505
Volume ID: VID00001
Source Institution: University of Florida
Holding Location: University of Florida
Rights Management: All rights reserved by the source institution and holding location.
Resource Identifier: ltqf - AAA0813
notis - ACK0935
alephbibnum - 000440469
oclc - 05069592
lccn - a 62009718

Full Text

'636- HUME iN


BULLET N AR

I.F.AS.- Utkof
OF THE


FLORIDA STATE MUSEUM

BIOLOGICAL SCIENCES


Volume 6


Number 2


OSTEOLOGY OF LIVING AND FOSSIL NEW WORLD
QUAILS (AVES, GALLIFORMES)

J. Alan Holman
09:


UNIVERSITY


OF FLORIDA


Gainesville
1961


Florlka


T III I . .


tp &G,







Numbers of the BULLETIN OF THE FLORIDA STATE MUSEUM are pub-
lished at irregular intervals. Volumes contain about 300 pages and are not
necessarily completed in any one calendar year.


o'6
/1





SCIENCE
LIBRARY


WILuIA J. RIEMER, Managing Editor
OLIVER L. AUSTIN, JR., Editor



Consultant for this issue:
Alexander Wetmore
frvffl---------------


3 1262 04279 7307












Communications concerning purchase or exchange of the publication and all
manuscripts should be addressed to the Managing Editor of the Bulletin, Florida
State Museum, Seagle Building, Gainesville, Florida.


Published 22 December 1961


Price for this issue $1.55









OSTEOLOGY OF LIVING AND FOSSIL NEW WORLD QUAILS
(AVES, GALLIFORMES)


J. ALAN HOLMAN







SYNOPSIS: The family Odontophoridae is divided into two distinct groups on the
basis of pelvic structure. The Dendrortyx group contains Miortyx of the Miocene,
Neortyx peninsularis new genus and species from the Pleistocene, and the living
genera Dendrortyx, Philortyx, Oreortyx, Callipepla, Colinus, and Lophortyx. Den-
drortyx is at once the most primitive and aberrant living genus of American quails
and is most closely related to Philortyx. Oreortyx is a near relative of the Calli-
pepla-Colinus-Lophortyx complex of genera. The members of this complex are
so similar that they might best be considered as subgenera.
The Odontophorus group contains the living genera Odontophorus, Dactylor-
tyx, Cyrtonyx, and Rhynchortyx. Odontophorus appears to be the most primitive
genus in this group. Dactylortyx and Cyrtonyx are similar. Rhynchortyx is the
most aberrant genus of the group.
The Miocene species Cyrtonyx tedfordi probab! 'Ivold be transferred from
the Odontophoridae to the Cracidae.
The study of more than 700 fossils of the genus Colinus shows a progressive
trend toward reduction in size and gradual change in qualitative characters.
The transition from C. hibbardi to C. suilium took place during the early Pleisto-
cene, and the change from C. suilium to C. virginianus occurred in Wicomico
time.








1The author is Associate Professor of Vertebrate Zoology at Illinois State Nor-
mal University, Normal. He has published several papers on amphibians, rep-
tiles, and mammals, living and fossil. One of these-Birds and Mammals from the
Pleistocene of Williston, Florida, 1959-appeared in this journal. The present
paper'represents in part a doctoral dissertation accepted at the University of Flor-
ida in 1961. Manuscript submitted 29 May 1961.-ED.


Holman, J. Alan. 1961. Osteology of living and fossil New World quails (Aves,
Galliformes). Bull. Florida State Mus., vol. 6, no. 2, pp. 131-233.


UNIVERSITY OF FLORIDA LIBRARIES






BULLETIN FLORIDA STATE MUSEUM


TABLE OF CONTENTS


Introduction ----------
Acknowledgments ---
Materials and methods ------

Osteology of living New
World quails -_


Dendrortyx -
Philortyx ---.
Oreortyx ------
Callipepla --
Colinus -------
Lophortyx ---
Odontophorus
Dactylortyx -
Cyrtonyx ---_
Rhynchortyx -
Summary of ge
Major relations


-- 132
133
------- 134


S134
-- 135
-- 138
---. 141


---- 145
-__ --- 148
..--......._........------ 153
-------158
* __.. . . . . . . 1 5 8
164
.--------__-__.---- 164
---------- 167
.------- 170
neric characters 173
hips ----_ 186


Miortyx


_------- 193


Cyrtonyx ----_ -- -
Colinus --
Lophortyx --------
[Generically undetermined


forms] --
Pleistocene forms --
Dendrortyx -
Oreortyx --
Neortyx, new ge
Colinus -----__


194
195
196


193 and 196
.----- 197
-- --- 197
-- --- 198
nus -------- 198
.--- 201


Lophortyx -----
Cyrtonyx ------
Odontophorus ---.-
Forms from archaeological
sites ---
Variation in fossil Colinus .


Conclusions --


- 203
204
204


-207


Fossil record of New World
quails _-__----- 193
Tertiary forms _---_------- 193


Tables


Literature cited


INTRODUCTION

The New World quails form a group of rather small-sized galliform
birds that range throughout the tropical, subtropical, and temperate
regions of the Americas. Most workers consider them a subfamily of
the Phasianidae, the Odontophorinae. The North American species
are well known individually, and a voluminous literature on such sub-
jects as life history, ecological tolerance, food and nutrition, and arti-
ficial propagation has grown out of their value as game birds. The
Central and South American forms are generally less well known.
Crispens (1960) lists literature on the economic and management as-
pects of the North American species. Ridgway and Friedmann (1946)
give an excellent bibliography on the taxonomy of North and Central
American forms; Hellmayr and Conover (1942) include the South
American forms.


Vol. 6


__






HOLMAN: OSTEOLOGY OF QUAILS


The classification of galliform taxa at the family level and above
is based mainly on internal anatomy, whereas that of taxa below the
family level is based on external features. Thus, the New World
quails are defined on the following characters (Ridgway and Fried-
mann, 1946): 1) mandibular tomium serrated or toothed; 2) tail less
than half as long to slightly longer than wing; 3) tarsus less than one-
fourth to more than one-third as long as wing; 4) bill relatively short
and stout; 5) rectrices 10 to 14; 6) tarsus without spurs; 7) acrotarsium
with a single row of broad, transverse scutellae; and 8) plant tarsi
with two or more definite rows of moderately long scutellae, but also
covered with small scales. The serrated or toothed mandibular to-
mium is unique in the order, whereas the other characters occur
singly or in various combinations in other families and subfamilies.
Genera of New World quails are defined on such characters as
proportionate lengths of bill and limbs, development of head plumes,
number of rectrices, relative length of primaries, markings of chest,
sides, and flanks, shape of tail, and scutellation of tarsus. Species and
subspecies are defined almost entirely on the basis of coloration,
markings, and size.
Most recent classifications place the New World quails with the
Old World quails and pheasants in the family Phasianidae (Ridgway
and Friedmann, 1946, and Wetmore, 1960), or the subfamily Phasiani-
nae (Mayr and Amadon, 1951, and Hudson, Lanzillotti, and Edwards,
1959). Sibley (1960) studied the electrophoretic properties of the egg-
white proteins of four genera of North American quails and tentatively
suggests that the New World quails appear to form as distinct a group
as certain other galliforms given family or subfamily rank in current
classifications. A study of the osteology of galliform birds by Holman
(MS) indicates that the New World quails represent a natural and
highly evolved group that should be recognized as a family, the Odon-
tophoridae, rather than as a subfamily of the Phasianidae.
No comprehensive comparative study has been published of the
internal anatomy of New World quails at the generic or infrageneric
level. The present work consists of two phases. The skeletons of
the living genera, species, and subspecies of New World quails, inso-
far as available, are described, and criteria for the various taxa are
established. The osteological variations of the fossil material are
evaluated.
ACKNOWLEDGMENTS
Deepest thanks go to Pierce Brodkorb for valuable advice and con-
stant encouragement throughout his supervision of this problem. Sin-







BULLETIN FLORIDA STATE MUSEUM


cere thanks are also due other biologists who critically read the manu-
script: Oliver L. Austin, Jr., Eugene Bovee, Harold K. Brooks, James
N. Layne, E. Lowe Pierce, and William J. Riemer. I am also grateful
to Ted T. Allen, who made the drawings, and Robert W. McFarlane,
who did the photographic work. To my wife, Betty Holman, who
has typed the material in this paper countless times, I owe special
thanks.
MATERIALS AND METHODS

The skeletal collection of Pierce Brodkorb at the University of Florida
was supplemented by material borrowed from the United States Na-
tional Museum (through Herbert Friedmann), University of Michigan
Museum of Zoology (through H. B. Tordoff), University of California
Museum of Zoology (through Frank A. Pitelka), and Southern Illinois
University Wildlife Research Collection (through W. D. Klimstra).
Fossil specimens studied in the present work are from the Brodkorb
collection at the University of Florida, the University of Oregon Mu-
seum of Natural History (through J. Arnold Shotwell), and the Texas
Memorial Museum (through John A. Wilson).
The number of specimens studied is listed below. Incomplete
skeletons are given in parentheses.
Dendrortyx leucophrys 1 C. leucopogon 4
Philortyx fasciatus 1 Lophortyx californica 2 (+ 1)
Oreortyx picta 4 L. gambelii 2
Callipepla squamata 4 L. douglasii 2
Colinus virginianus floridanus 9 (+ 1) Odontophorus guianensis 1 (+ 1)
C. v. virginianus x C. v. floridanus 23 0. stellatus 1
C. v. insignis 2 (+ 3) 0. guttatus 1
C. v. coyolcos (1) Dactylortyx thoracicus 3
C. v. virginianus 65 Cyrtonyx montezumae 3
C. nigrogularis (2) Rhynchortyx cinctus 1

The fossil specimens include 795 elements, detailed in the body of
the work.
All measurements are linear and to the nearest tenth millimeter.
Although 60 intramembral and intermembral ratios were taken, only
those that proved useful are presented herein.
Anatomical nomenclature follows that of Howard (1929).

OSTEOLOGY OF LIVING NEW WORLD QUAILS
This section presents an osteological account of the living genera and
species of the New World quails available for study and remarks on
skeletal variation in some subspecies of Colinus virginianus. Charac-


Vol 6






HOLMAN: OSTEOLOGY OF QUAILS


ters at the generic level are numbered consecutively to facilitate in-
tergeneric comparisons by the reader. The skeletal elements studied
are those bones most frequently found as fossils.

Dendrortyx GOULD, 1844
This genus includes four living species, confined to the mountains of
Mexico and Central America. Only a single skeleton of Dendrortyx
leucophrys (Gould, 1844) was available. Its measurements are given
in table 1.
ROSTRUM. (1) Relatively long and shallow. (2) Nasal fossae ellip-
tical.
STERNUM. (3) Height of carina through sternal plate 30 percent
length of sternum. (4) Dorsal lip of coracoidal sulcus projected well
beyond ventral lip. (5) Depressions on anterior portion of sternal
plate shallow and slightly pneumatic. (6) Base of posterior lateral pro-
cess more than one-half width of sternal plate at its middle.
CoRAcom. (7) Entire proximal end bending ventrad above level
of procoracoid process. (8) Head broadly rounded, much depressed,
and much inflected. (9) Brachial tuberosity with moderate dorsal, but
slight ventral overhang. (10) Furcular facet weakly inflated. (11)
Bicipital attachment well developed; margin between it and head
concave. (12) Shaft narrow proximally, with distal portion of its ven-
tral surface convex medially. (13) Procoracoid process with its apex
rounded. (14) Dorsal intermuscular line swinging out to lateral border
of shaft; distal fifth sharply raised. (15) Ventral intermuscular line
terminating in middle of distal border of sterno-coracoidal process.
(16) Dorsal face weakly excavated distally, but with distinct round,
deep, fossa below tubercle for ligamentum sterno-coracoideum dorsale,
and above external end of sternal facet. (17) Tubercle for ligamentum
sterno-coracoideum dorsale obsolete, but rounded. (18) Internal distal
angle without distinct tubercle or ridge on dorsal face. (19) Sterno-
coracoidal process with distal margin strongly oblique, not produced
as a sharply raised ridge, but grooved dorsally, its tip directed prox-
imally, but without a terminal knob. (20) Distal border of sternal
facet shallowly concave.
SCAPULA. (21) Glenoid facet elliptical in dorsal view. (22) Dorsal
depression just media to glenoid facet deeply excavated. (23) Bridge
between acromion process and glenoid facet about one-half width of
glenoid facet. (24) Acromion process with its apex acute and slightly
deflected.






BULLETIN FLORIDA STATE MUSEUM


HUMERUS. (25) Pneumatic fossa a short oval, without inner shelf.
(26) Median crest strongly developed, elevated throughout. (27)
Capital groove deeply excavated, its internal margin strongly elevated.
(28) Margin between head and internal tuberosity only slightly con-
cave. (29) Head about as long as broad in anconal view. (30) Ridge
along medial border of fossa II slightly swollen, elongate, and with
its apex acute. (31) Fossa II obsolete. (32) Latissimus ridge (Ashley,
1941) extending along lateral edge of shaft throughout its length.
(33) External tuberosity moderately inflated. (34) Bicipital crest aris-
ing gently from shaft, rounded, and with its distal border lacking
a groove. (35) Deltoid crest strongly developed, much inflected, and
its summit knoblike. (36) Entepicondyle at level of internal condyle.
(37) Entepicondylar prominence weakly produced, its pit placed on
internal face. (38) Depression of M. brachialis anticus deeply exca-
vated. (39) Distal border of external condyle flat in anconal view.
ULNA. (40) Olecranon process strongly produced, its apex round-
ed. (41) External cotyla shortened, flattened dorsally, its distal border
sloping precipitously into shaft and deeply notched by proximal radial
impression. (42) Impression for M. brachialis anticus excavated, its
borders distinct. (43) Height at middle of shaft 83 percent of height
just distal to external cotyla (44) Distal radial impression well exca-
vated. (45) Notch between carpal tuberosity and internal condyle
weak. (46) External condyle weakly produced, arising abruptly from
shaft, and its ventral border flattened.
CARPOMETACARPUS. (47) Metacarpal I moderately pointed ter-
minally. (48) Ridge separating anterior carpal fossa from excavated
area above pisiform process weak and extending only slightly onto
base of metacarpal I. (49) Pisiform process round in internal view,
much reflected dorsad, thus produced well above dorsal rim of inner
carpal trochlea. (50) Ligamental attachment of pisiform process pro-
duced throughout, moderately swollen proximally and distally, with
proximal portion separated from pisiform process by shallow sulcus.
(51) Internal carpal trochlea with its proximal border shallowly con-
cave, its ventral border slightly notched. (52) Intermetacarpal tu-
bercle extending to and ankylosing with metacarpal III, and distally
placed with a small space between its posterior border and junction
of metacarpals II and III. (53) Tubercle on internal proximal face of
metacarpal III obsolete. (54) One moderately developed and one
obsolete tubercle on external proximal face of metacarpal III. (55)
Metacarpal III moderately bowed.
PELVIS. (56) Width of pelvis through antitrochanters 51 percent
length of innominate from anterior edge of ilium to ischial angle. (57)


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HOLMAN: OSTEOLOGY OF QUAILS


Pectineal process obsolete, only moderately produced beyond acetab-
ular border, and its apex rounded. (58) Depression on medial side of
pectineal process deeply excavated and perforated by a foramen.
(59) Posterior iliac crest without prominence. (60) Ridge originating
on posterior iliac crest at level of ilio-ischiac fenestra and continuing
ventrad and posteriad toward posterior border of ilium with prom-
inence arising sharply from its posterior portion. (61) Dorsal face of
postacetabular ilium moderately broad anteriad, narrowing abruptly
posteriad to form acute apex of short, broad, dorsal roof of posterior
process. (62) Caudal face of postacetabular ilium enlarged, triangu-
lar, and vertical in position, its posterodorsal apex forming posterome-
dial wall of posterior process. (63) Lateral face of postacetabular
ilium with its posterodorsal apex acute and forming shortened lateral
wall of posterior process. (64) Renal bar slender; both bar and poste-
rior process excavated by renal depression. (65) Ischium at angle of
about 160 degrees to ventral edge of posterior iliac crest. (66) Width
through transverse processes of fourth synsacral vertebra 55 percent
of length of synsacrum from centrum of first synsacral vertebra through
transverse process of fourth synsacral vertebra.
FEMUR. (67) Distal width 17 percent of length. (68) Head mod-
erately enlarged, bent dorsally, and rotated posteriorly. (69) Neck
long. (70) Iliac facet compressed; posterior border straight. (71)
Dorsal crest of trochanter depressed and deflected, its dorsal border
flat. (72) Trochanteric ridge 20 percent length of femur. (73) Distal
depth 85 percent of distal width. (74) Internal condyle moderately
wide, its anterior border much produced throughout. (75) External
condyle depressed, flattened anteriorly.
TIBIOTARSUS. (76) Depth of inner cnemial crest 59 percent of its
length. (77) Outer cnemial crest decurved. (78) Interarticular tu-
bercle moderately produced. (79) Area between external articular
surface and outer cnemial crest moderately concave in dorsal view.
(80) Posterior margin between external and internal articular surfaces
weakly notched. (81) Area just anterior to ridge from proximal in-
ternal ligamental attachment with deeply excavated depression. (82)
Internal ligamental prominence on internal condyle strongly pro-
duced. (83) Face of internal condyle well excavated, its border highly
elevated. (84) Dorsal border of supratendinal bridge oblique.
TARSOMETATARSUS. (85) Distal width 14 percent of length. (86)
Proximal depth 94 percent of proximal width. (87) Intercotylar prom-
inence moderately deflected, its apex pointed. (88) Posterior inter-
cotylar area moderately excavated as a triangular depression. (89)
Area surrounding posterior opening of inner proximal foramen deeply


1961






BULLETIN FLORIDA STATE MUSEUM


excavated. (90) Hypotarsus with 2 well-developed and 1 obsolete
calcaneal ridges, 2 calcaneal grooves, and 1 closed calcaneal canal.
(91) Anterior face of shaft extensively excavated by anterior metatarsal
groove. (92) Depth of middle trochlea 56 percent of distal width. (93)
Trochlea for digit II only slightly inflected. (94) Wing of trochlea for
digit IV depressed anteroposteriorly; thus entire trochlea slightly long-
er than deep.
INTERMEMBRAL PROPORTIONS. (95) Wing/leg 52 percent. (96)
Ulna/humerus 88 percent. (97) Tarsometatarsus/femur 88 percent.
(98) Tarsometatarsus/tibiotarsus 63 percent. (99) Humerus/femur 71
percent. (100) Humerus/tibiotarsus 51 percent. (101) Ulna/femur
63 percent. (102) Ulna/tibiotarsus 45 percent. (103) Ulna/tarsometa-
tarsus 72 percent. (104) Carpometacarpus/femur 37 percent. (105)
Carpometacarpus/tibiotarsus 27 percent. (106) Carpometacarpus/tar-
sometatarsus 42 percent. (107) Tarsometatarsus/humerus 124 percent.
(108) Ilium/sternum 89 percent. (109) Height of sternal carina/ilium
34 percent.
Philortyx GOULD, 1846
This monotypic genus is confined to the mountains of Mexico. Meas-
urements of Philortyx fasciatus (Gould, 1844) are given in table 1.
ROSTRUM. (1) Relatively long and shallow. (2) Nasal fossae ellip-
tical.
STERNUM. (3) Sternum incomplete. (4) Dorsal lip of coracoidal
sulcus projected well beyond ventral lip. (5) Depressions on anterior
portion of sternal plate deep and slightly pneumatic. (6) Base of
posterior lateral process less than one-half width of sternal plate at
its middle.
CORACOID. (7) Entire proximal end straight above procoracoid
process. (8) Head pointed, much compressed, and much reflected.
(9) Brachial tuberosity obsolete dorsally, but with moderate ventral
overhang. (10) Furcular facet weakly inflated. (11) Bicipital attach-
ment well developed; margin between it and head concave. (12) Shaft
wide proximally, distal portion of its ventral surface concave medially.
(13) Procoracoid process with its apex acute. (14) Dorsal intermuscu-
lar line swinging out to medial surface of shaft, distal half sharply
raised. (15) Ventral intermuscular line terminating at tip of sterno-
coracoidal process. (16) Dorsal face weakly excavated distally, with-
out fossa. (17) Tubercle for ligamentum sterno-coracoideum dorsale
well developed and rounded. (18) Internal distal angle with distinct
dorsal ridge extending obliquely to above lateral end of sternal facet.
(19) Sterno-coracoidal process with distal margin moderately oblique,


Vol. 6






HOLMAN: OSTEOLOGY OF QUAILS


not produced as a sharply raised ridge, and without dorsal groove;
its tip directed laterally and ending in an obsolete terminal knob. (20)
Distal border of sternal facet shallowly concave.
SCAPULA. (21) Glenoid facet oval in dorsal view. (22) Dorsal de-
pression just media to glenoid facet shallowly excavated. (23) Bridge
between acromion process and glenoid facet about one-third width of
glenoid facet. (24) Acromion process with its apex rounded and mod-
erately deflected.
HUMERUS. (25) Pneumatic fossa a moderately long ellipse with-
out inner shelf. (26) Median crest weakly developed, but elevated
throughout. (27) Capital groove deeply excavated, its internal mar-
gin strongly elevated. (28) Margin between head and internal tuberos-
ity moderately concave. (29) Head about as long as broad in anconal
view. (30) Ridge along medial border of fossa II slightly swollen,
elongate, its apex acute. (31) Fossa II well developed. (32) Latissi-
mus ridge originating on lateral edge of shaft, but swinging in onto
anconal face proximally. (33) External tuberosity moderately inflated.
(34) Bicipital crest arising gently from shaft, pointed, its distal border
lacking a groove. (35) Deltoid crest moderately developed, slightly
inflected, its summit knoblike. (36) Entepicondyle produced slightly
beyond level of internal condyle. (37) Entepicondylar prominence
moderately produced, its pit placed on internal face. (38) Depression
for M. brachialis anticus deeply excavated. (39) Distal border of ex-
ternal condyle rounded in anconal view.
ULNA. (40) Olecranon process strongly produced, its apex pointed.
(41) External cotyla moderately elongate, rounded throughout, its dis-
tal border sloping gently into shaft and deeply notched by proximal
radial impression. (42) Impression for M. brachialis anticus repre-
sented by an unexcavated scar, its borders indistinct. (43) Height at
middle of shaft 85 percent of height just distal to external cotyla. (44)
Distal radial impression obsolete. (45) Notch between carpal tuberos-
ity and internal condyle well developed. (46) External condyle well
produced, arising abruptly from shaft, its ventral border flattened.
CARPOMETACARPUS. (47) Metacarpal I moderately pointed termi-
nally. (48) Ridge separating anterior carpal fossa from excavated area
above pisiform process produced and extending well onto base of
metacarpal I. (49) Pisiform process round in internal view, moderately
reflected dorsad, thus produced slightly above dorsal rim of inner car-
pal trochlea. (50) Ligamental attachment of pisiform process pro-
duced and moderately swollen proximally, flattened distally, with prox-
imal portion connected to pisiform process by bony bridge. (51) In-
ternal carpal trochlea with its proximal border shallowly concave, its







BULLETIN FLORIDA STATE MUSEUM


ventral border without notch. (52) Intermetacarpal tubercle extending
slightly beyond, but not ankylosing with metacarpal III, distally lo-
cated with a small space between its posterior border and junction of
metacarpals II and III. (53) Tubercle on internal proximal face of
metacarpal III elongate and weakly produced. (54) Only one mod-
erately developed tubercle on external proximal face of metacarpal
III. (55) Metacarpal III slightly bowed.
PELVIS. (56) Width of pelvis through antitrochanters 47 percent
length of innominate from anterior edge of ilium to ischial angle.
(57) Pectineal process obsolete, only moderately produced beyond
acetabular border, its apex pointed. (58) Depression on medial side
of pectineal process deeply excavated, but without foramen. (59)
Posterior iliac crest without prominence. (60) Ridge originating on
posterior iliac crest at level of ilio-ischiac fenestra, continuing ventrad
and posteriad towards posterior border of. ilium with prominence
sharply arising from its posterior portion. (61) Dorsal face of post-
acetabular ilium moderately broad anteriad, narrowing abruptly pos-
teriad to form acute apex of short, broad, dorsal roof of posterior
process. (62) Caudal face of postacetabular ilium enlarged, triangular,
and vertical in position, its posterodorsal apex forming posteromedial
wall of posterior process. (63) Lateral face of postacetabular ilium
with its posterodorsal apex acute and forming shortened lateral wall
of posterior process. (64) Renal bar moderately slender; both bar and
posterior process excavated by renal depression. (65) Ischium at angle
of about 160 degrees to ventral edge of posterior iliac crest. (66)
Width through transverse process of fourth synsacral vertebra 60 per-
cent of length of synsacrum from centrum of first synsacral vertebra
through transverse process of fourth synsacral vertebra.
FEMUR. (67) Distal width 14 percent of length. (68) Head mod-
erately enlarged, neither bent dorsally nor rotated posteriorly. (69)
Neck only moderately long. (70) Iliac facet without compression,
posterior border convex. (71) Dorsal crest of trochanter compressed
and reflected, its dorsal border rounded. (72) Trochanteric ridge 27
percent length of femur. (73) Distal depth 97 percent of distal width.
(74) Internal condyle wide, its anterior border moderately produced
throughout. (75) External condyle compressed and rounded anteri-
orly.
TIBIOTARSUS. (76) Depth of inner cnemial crest about 45 percent
of its length. (77) Outer cnemial crest decurved. (78) Interarticular
tubercle weakly produced. (79) Area between external articular sur-
face and outer cnemial crest moderately concave in dorsal view. (80)
Posterior margin between external and internal articular surface with


Vol. 6






HOLMAN: OSTEOLOGY OF QUAILS


a distinct notch. (81) Area just anterior to ridge from proximal in-
ternal ligamental attachment without deeply excavated depression.
(82) Internal ligamental prominence on internal condyle moderately
produced. (83) Face of internal condyle moderately excavated, its
border moderately elevated. (84) Dorsal border of supratendinal
bridge straight.
TARSOMETATARSUS. (85) Distal width 17 percent of length. (86)
Proximal depth 86 percent of proximal width. (87) Intercotylar prom-
inence much deflected, its apex pointed. (88) Posterior intercotylar
area weakly excavated as a triangular depression. (89) Area surround-
ing posterior opening of inner proximal foramen deeply excavated.
(90) Hypotarsus with 2 well-developed and 1 obsolete calcaneal ridges,
2 calcaneal grooves, and 2 closed calcaneal canals. (91) Anterior face
of shaft extensively excavated by anterior metatarsal groove. (92)
Depth of middle trochlea 59 percent of distal width. (93) Trochlea
for digit II only slightly inflected. (94) Wing of trochlea for digit IV
depressed dorsoventrally, thus entire trochlea much deeper than long.
INTERMEMBRAL PROPORTIONS. (95) Wing/leg 65 percent. (96)
Ulna/humerus 87 percent. (97) Tarsometatarsus/femur 76 percent.
(98) Tarsometatarsus/tibiotarsus 59 percent. (99) Humerus/femur 82
percent. (100) Humerus/tibiotarsus 64 percent. (101) Ulna/femur 72
percent. (102) Ulna/tibiotarsus 55 percent. (103) Ulna/tarsometa-
tarsus 94 percent. (104) Carpometacarpus/femur 44 percent. (105)
Carpometacarpus/tibiotarsus 34 percent. (106) Carpometacarpus/tar-
sometatarsus 58 percent. (107) Tarsometatarsus/humerus 93 percent.
(108) Ilium/sternum 41 percent. (109) Height of sternal carina/ilium
41 percent.
Oreortyx BAIRD, 1858
The monotypic genus Oreortyx occurs from Washington to northern
Baja California. Measurements of 0. picta (Douglas, 1829) are given
in table 1.
ROsTRUM. (1) Relatively long and shallow. (2) Nasal fossae ellipti-
cal.
STERNUM. (3) Height of carina through sternal plate 29 to 31
percent length of sternum. (4) Dorsal lip of coracoidal sulcus pro-
jected well beyond ventral lip. (5) Depressions on anterior portion
of sternal plate deep and moderately pneumatic. (6) Base of posterior
lateral process less than one-half width of sternal plate at its middle.
CORACOID. (7) Entire proximal end straight above procoracoid
process. (8) Head broadly rounded, moderately depressed, and mod-
erately reflected. (9) Brachial tuberosity with slight dorsal and ventral






BULLETIN FLORIDA STATE MUSEUM


overhang. (10) Furcular facet moderately inflated. (11) Bicipital at-
tachment well developed, margin between it and head concave. (12)
Shaft wide proximally, distal portion of its ventral surface concave
medially. (13) Procoracoid process with its apex rounded. (14) Dor-
sal intermuscular line swinging out to medial surface of shaft, distal
half sharply raised. (15) Ventral intermuscular line terminating at tip
of sterno-coracoidal process. (16) Dorsal face weakly excavated dis-
tally, without fossa. (17) Tubercle for ligamentum sterno-coracoideum
dorsale well developed, but irregular in outline. (18) Internal distal
angle with distinct irregular tubercle extending obliquely to above
lateral end of sternal facet. (19) Sterno-coracoidal process with distal
margin slightly oblique, not produced as a sharply raised ridge, and
without dorsal groove; its tip directed laterally and ending in a termi-
nal knob. (20) Distal border of sternal facet deeply concave.
SCAPULA. (21) Glenoid facet oval in dorsal view. (22) Dorsal de-
pression just media to glenoid facet shallowly to moderately exca-
vated. (28) Bridge between acromion process and glenoid facet about
one-half width of glenoid facet. (24) Acromion process with its apex
acute and slightly deflected.
HUMERUS. (25) Pneumatic fossa a moderately long oval or oval-
oid, without inner shelf. (26) Median crest strongly developed, ele-
vated only at its middle. (27) Capital groove moderately excavated,
its internal margin moderately elevated. (28) Margin between head
and internal tuberosity moderately concave. (29) Head broader than
long in anconal view. (30) Ridge along medial border of fossa II much
swollen, elongate, its apex acute. (31) Fossa II well developed. (32)
Latissimus ridge originating on lateral edge of shaft, but swinging in
onto anconal face proximally. (33) External tuberosity much inflated.
(34) Bicipital crest arising gently from shaft, pointed, its distal border
lacking a groove. (35) Deltoid crest moderately developed, slightly
inflected, its summit knoblike. (36) Entepicondyle at level of internal
condyle. (37) Entepicondylar prominence moderately produced, its
pit placed on internal face. (38) Depression of M. brachialis antics
shallowly excavated. (39) Distal border of external condyle rounded
in anconal view.
ULNA. (40) Olecranon process strongly produced, its apex pointed.
(41) External cotyla much elongated, rounded throughout, its distal
border sloping gently into shaft and only slightly notched by proximal
radial impression. (42) Impression for M. brachialis anticus excavated,
its borders distinct. (43) Height at middle of shaft 64 to 70 percent
of height just distal to external cotyla. (44) Distal radial impression
obsolete. (45) Notch between carpal tuberosity and internal condyle


Vol. 6






HOLMAN: OSTEOLOGY OF QUAILS


well developed. (46) External condyle moderately produced, arising
gently from shaft, rounded throughout.
CARPOMETACARPUS. (47) Metacarpal I sharply pointed terminally.
(48) Ridge separating anterior carpal fossa from excavated area above
pisiform process produced, and extending well onto base of metacar-
pal I. (49) Pisiform process ovaloid in internal view, slightly reflected
dorsad, thus below level of dorsal rim of inner carpal trochlea. (50)
Ligamental attachment of pisiform process produced and much swol-
len proximally, flattened distally, with proximal portion connected to
pisiform process by bony bridge. (51) Internal carpal trochlea with
proximal border distinctly concave and ventral border slightly notched.
(52) Intermetacarpal tubercle extending slightly beyond, but not an-
kylosing with metacarpal III, distally placed with a small space be-
tween its posterior border and junction of metacarpals II and III.
(53) Tubercle on internal proximal face of metacarpal III elongate and
moderately produced. (54) One strongly developed tubercle on ex-
ternal proximal face of metacarpal III. (55) Metacarpal III slightly
bowed.
PELVIS. (56) Width of pelvis through antitrochanters 60 to 63 per-
cent of length of innominate from anterior edge of ilium to ischial
angle. (57) Pectineal process obsolete, only moderately produced be-
yond acetabular border, its apex pointed. (58) Depression on medial
side of pectineal process shallowly excavated, without foramen. (59)
Posterior iliac crest without prominence. (60) Ridge originating on
posterior iliac crest at level of ilio-ischiac fenestra, and continuing ven-
trad and posteriad toward posterior border of ilium with prominence
weakly arising from its posterior portion. (61) Dorsal face of post-
acetabular ilium much broadened anteriad, narrowing abruptly pos-
teriad to form acute apex of short, broad, dorsal roof of posterior
process. (62) Caudal face of postacetabular ilium enlarged, broadly
triangular, vertical in position, its posterodorsal apex forming postero-
medial wall of posterior process. (63) Lateral face of postacetabular
ilium with posterodorsal apex acute, forming shortened lateral wall
of posterior process. (64) Renal bar obsolete, both bar and posterior
process excavated by renal depression. (65) Ischium at angle of
about 160 degrees to ventral edge of posterior iliac crest. (66) Width
through transverse process of fourth synsacral vertebra 67 to 85 per-
cent length of synsacrum from centrum of first synsacral vertebra
through transverse process of fourth synsacral vertebra.
FEMUR. (67) Distal width about 17 percent length. (68) Head
moderately enlarged, neither bent dorsally nor rotated posteriorly.
(69) Neck only moderately long. (70) Iliac facet without compression,






BULLETIN FLORIDA STATE MUSEUM


posterior border convex. (71) Dorsal crest of trochanter compressed
and reflected, its dorsal border rounded. (72) Trochanteric ridge 23
to 25 percent length of femur. (73) Distal depth 76 to 83 percent of
distal width. (74) Internal condyle moderately wide, its anterior bor-
der much produced throughout. (75) External condyle compressed,
rounded anteriorly.
TIBIOTARSUS. (76) Depth of inner cnemial crest 65 to 74 percent
of its length. (77) Outer cnemial crest straight. (78) Interarticular
tubercle strongly produced. (79) Area between external articular sur-
face and outer cnemial crest moderately concave in dorsal view. (80)
Posterior margin between external and internal articular surface with
a distinct notch. (81) Area just anterior to ridge from proximal internal
ligamental attachment without deeply excavated depression. (82)
Internal ligamental prominence on internal condyle moderately pro-
duced. (83) Face of internal condyle moderately excavated, its border
moderately elevated. (84) Dorsal border of supratendinal bridge usu-
ally straight.
TARSOMETATARSUS. (85) Distal width 17 to 18 percent of length.
(86) Proximal depth 91 to 98 percent of proximal width. (87) Inter-
cotylar prominence moderately deflected, its apex usually pointed.
(88) Posterior intercotylar area usually unexcavated, but may be
weakly excavated as a triangular depression. (89) Area surrounding
posterior opening of inner proximal foramen shallowly excavated.
(90) Hypotarsus with 2 well-developed and 1 obsolete calcaneal ridges,
2 calcaneal grooves, and usually 1, sometimes 2, closed calcaneal
canals. (91) Anterior metatarsal groove reduced or absent. (92) Depth
of middle trochlea 50 to 53 percent of distal width. (93) Trochlea for
digit II moderately inflected. (94) Wing of trochlea for digit IV de-
pressed dorsoventrally, thus entire trochlea much deeper than long.
INTERMEMBRAL PROPORTIONS. (95) Wing/leg 64 to 67 percent.
(96) Ulna/humerus 92 to 94 percent. (97) Tarsometatarsus/femur 75
to 82 percent. (98) Tarsometatarsus/tibiotarsus 58 percent. (99) Hu-
merus/femur 83 to 89 percent. (100) Humerus/tibiotarsus 58 to 61
percent. (101) Ulna/femur 78 to 82 percent. (102) Ulna/tibiotarsus
55 to 58 percent. (103) Ulna/tarsometatarsus 95 to 110 percent. (104)
Carpometacarpus/femur 47 to 50 percent. (105) Carpometacarpus/
tibiotarsus 33 to 35 percent. (106) Carpometacarpus/tarsometatarsus
57 to 66 percent. (107) Tarsometatarsus/humerus 85 to 99 percent.
(108) Ilium/sternum 65 to 69 percent. (109) Height of sternal carina/
ilium 45 to 46 percent.


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HOLMAN: OSTEOLOGY OF QUAILS


Callipepla WAGLER, 1832
The single species, Callipepla squamata (Vigors, 1830) occurs in the
southwestern United States and northern Mexico. Measurements are
given in table 1.
ROSTRUM. (1) Relatively long and shallow. (2) Nasal fossae ellipti-
cal.
STERNUM. (3) Height of carina through sternal plate 30 to 31 per-
cent length of sternum. (4) Dorsal lip of coracoidal sulcus projected
well beyond ventral lip. (5) Depressions on anterior portion of sternal
plate obsolete to deep, slightly to moderately pneumatic. (6) Base
of posterior lateral process less than one-half width of sternal plate at
its middle.
CORACOID. (7) Entire proximal end straight above procoracoid
process. (8) Head narrowly rounded, moderately compressed, mod-
erately inflected. (9) Brachial tuberosity with slight dorsal and ventral
overhang. (10) Furcular facet moderately inflated. (11) Bicipital at-
tachment well developed, margin between it and head concave. (12)
Shaft wide proximally, distal portion of its ventral surface usually con-
cave medially. (13) Procoracoid process with apex acute. (14) Dorsal
intermuscular line swinging out to medial surface of shaft, distal half
sharply raised. (15) Ventral intermuscular line terminating at tip of
sterno-coracoidal process. (16) Dorsal face weakly excavated distally,
without fossa. (17) Tubercle for ligamentum sterno-coracoideum dor-
sale obsolete, but rounded. (18) Internal distal angle with distinct
irregular tubercle extending obliquely to above lateral end of sternal
facet. (19) Sterno-coracoidal process with distal margin slightly
oblique, not produced as a sharply raised ridge, without dorsal groove;
its tip directed laterally and ending in a terminal knob. (20) Distal bor-
der of sternal facet usually deeply concave.
SCAPULA. (21) Glenoid facet oval in dorsal view. (22) Dorsal de-
pression just media to glenoid facet unexcavated, shallowly exca-
vated, or moderately excavated. (23) Bridge between acromion process
and glenoid facet about one-half width of glenoid facet. (24) Acrom-
ion process with its apex rounded, slightly deflected.
HUMERUS. (25) Pneumatic fossa a moderately long oval or ovaloid,
without inner shelf. (26) Median crest usually weakly developed, ele-
vated only at its middle. (27) Capital groove moderately excavated,
its internal margin moderately elevated. (28) Margin between head
and internal tuberosity moderately concave. (29) Head broader than
long in anconal view. (30) Ridge along medial border of fossa II mod-
erately swollen, elongate, its apex acute. (31) Fossa II well developed.






BULLETIN FLORIDA STATE MUSEUM


(32) Latissimus ridge originating on lateral edge of shaft, but swinging
in onto anconal face proximally. (33) External tuberosity moderately
inflated. (34) Bicipital crest arising gently from shaft, narrowly round-
ed, its distal border lacking a groove. (35) Deltoid crest moderately
developed, slightly inflected, its summit knoblike. (36) Entepicondyle
at level of internal condyle. (37) Entepicondylar prominence moder-
ately produced, its pit on palmar face. (38) Depression of M. brachi-
alis anticus usually shallowly excavated. (39) Distal border of external
condyle rounded in anconal view.
ULNA. (40) Olecranon process strongly produced, its apex pointed.
(41) External cotyla moderately elongate, rounded throughout, its dis-
tal border sloping gently into shaft, only slightly notched by proximal
radial impression. (42) Impression for M. brachialis anticus excavated,
its borders distinct. (43) Height at middle of shaft 63 to 66 percent
of height just distal to external cotyla. (44) Distal radial impression
obsolete. (45) Notch between carpal tuberosity and internal condyle
well developed. (46) External condyle well produced, arising abruptly
from shaft, rounded throughout.
CARPOMETACARPUS. (47) Metacarpal I moderately pointed termi-
nally. (48) Ridge separating anterior carpal fossa from excavated area
above pisiform process produced, extending well onto base of meta-
carpal I. (49) Pisiform process round or ovaloid in internal view,
slightly reflected dorsad, thus usually below, occasionally at level of
dorsal rim of inner carpal trochlea. (50) Ligamental attachment of
pisiform process produced and much swollen proximally, flattened dis-
tally, with proximal portion connected to pisiform process by bony
bridge. (51) Internal carpal trochlea with proximal border deeply
notched, its ventral border without notch. (52) Intermetacarpal tu-
bercle extending to, but not ankylosing with metacarpal III, distally
placed with a small space between its posterior border and junction
of metacarpals II and III. (53) Tubercle on internal proximal face
of metacarpal III obsolete. (54) One strongly developed tubercle on
external proximal face of metacarpal III. (55) Metacarpal III slightly
bowed.
PELVIS. (56) Width of pelvis through antitrochanters 58 to 62 per-
cent of length of innominate from anterior edge of ilium to ischial
angle. (57) Pectineal process obsolete, only moderately produced be-
yond acetabular border, its apex pointed. (58) Depression on medial
side of pectineal process shallowly excavated, without foramen. (59)
Posterior iliac crest without prominence. (60) Ridge originating on
posterior iliac crest at level of ilio-ischiac fenestra, continuing ventrad
and posteriad toward posterior border of ilium with prominence


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HOLMAN:- OSTEOLOGY OF QUAILS


weakly arising from its posterior portion. (61) Dorsal face of post-
acetabular ilium moderately broad anteriad, narrowing abruptly pos-
teriad to form acute apex of short, broad, dorsal roof of posterior
process. (62) Caudal face of postacetabular ilium enlarged, triangular,
vertical in position, its posterodorsal apex forming posteromedial wall
of posterior process. (63) Lateral face of postacetabular ilium with
its posterodorsal apex acute, forming shortened lateral wall of pos-
terior process. (64) Renal bar slender, both bar and posterior process
excavated by renal depression. (65) Ischium at angle of about 160
degrees to ventral edge of posterior iliac crest. (66) Width through
transverse process of fourth synsacral vertebra 71 to 80 percent of
length of synsacrum from centrum of first synsacral vertebra through
transverse process of fourth synsacral vertebra.
FEMUR. (67) Distal width 17 to 18 percent of length. (68) Head
moderately enlarged, neither bent dorsally nor rotated posteriorly.
(69) Neck only moderately long. (70) Iliac facet without compression,
posterior border convex. (71) Dorsal crest of trochanter compressed
and reflected, its dorsal border rounded. (72) Trochanter ridge 24 per-
cent length of femur. (73) Distal depth 76 to 82 percent of distal
width. (74) Internal condyle moderately wide, its anterior border much
produced throughout. (75) External condyle compressed and rounded
anteriorly.
TIBIOTARSUS. (76) Depth of inner cnemial crest 64 to 73 percent
of its length. (77) Outer cnemial crest usually straight. (78) Inter-
articular tubercle moderately produced. (79) Area between external
articular surface and outer cnemial crest moderately concave in dor-
sal view. (80) Posterior margin between external and internal articular
surfaces with distinct notch. (81) Area just anterior to ridge from proxi-
mal internal ligamental attachment without deeply excavated depres-
sion. (82) Internal ligamental prominence on internal condyle mod-
erately produced. (83) Face of internal condyle moderately exca-
vated, its border moderately elevated. (84) Dorsal border of supra-
tendinal bridge straight.
TARSOMETATARSUS. (85) Distal width 15 to 18 percent of length.
(86) Proximal depth 95 to 98 percent of proximal width. (87) Inter-
cotylar prominence moderately deflected, its apex usually pointed.
(88) Posterior intercotylar area weakly excavated as a triangular de-
pression. (89) Area surrounding posterior opening of inner proximal
foramen shallowly excavated. (90) Hypotarsus with 2 well-developed
and 1 obsolete calcaneal ridges, 2 calcaneal grooves, and 2 closed cal-
caneal canals. (91) Anterior face of shaft moderately excavated by






BULLETIN FLORIDA STATE MUSEUM


anterior metatarsal groove. (92) Depth of middle trochlea 52 to 59
percent of distal width. (93) Trochlea for digit II only slightly in-
flected. (94) Wing of trochlea for digit IV depressed dorsoventrally,
thus entire trochlea much deeper than long.
INTERMEMBRAL PROPORTIONS. (95) Wing/leg 67 to 69 percent. (96)
Ulna/humerus 92 percent. (97) Tarsometatarsus/femur 79 to 82 per-
cent. (98) Tarsometatarsus/tibiotarsus 59 percent. (99) Humerus/
femur 85 to 89 percent. (100) Humerus/tibiotarsus 63 to 65 percent.
(101) Ulna/femur 78 to 82 percent. (102) Ulna/tibiotarsus 58 to 59
percent. (103) Ulna/tarsometatarsus 95 to 102 percent. (104) Carpo-
metacarpus/femur 47 to 48 percent. (105) Carpometacarpus/tibio-
tarsus 34 to 35 percent. (106) Carpometacarpus/tarsometatarsus 58 to
61 percent. (107) Tarsometatarsus/humerus 89 to 96 percent. (108)
Ilium/sternum 66 to 69 percent. (109) Height of sternal carina/ilium
43 to 47 percent.
Colinus GOLDFUSS, 1820
The genus Colinus occurs in the United States east of the Rockies, on
Cuba and the Isle of Pines, and through Mexico and Central America
to Colombia and Venezuela. Of the four living species, table 1 in-
cludes measurements of C. virginianus (Linnaeus, 1758), C. nigrogul-
aris (Gould, 1843), and C. leucopogon (Lesson, 1842).
RosTRUM. (1) Relatively long and shallow. (2) Nasal fossae ellipti-
cal.
STERNUM. (3) Height of carina through sternal plate 25 to 32 per-
cent length of sternum. (4) Dorsal lip of coracoidal sulcus projected
well beyond ventral lip. (5) Depressions on anterior portion of sternal
plate usually shallow and nonpneumatic. (6) Base of posterior lateral
process less than one-half width of sternal plate at its middle.
CORACom. (7) Entire proximal end straight above procoracoid
process. (8) Head narrowly rounded, moderately compressed, mod-
erately reflected or inflected. (9) Brachial tuberosity with slight dor-
sal and ventral overhang. (10) Furcular facet weakly or moderately
inflated. (11) Bicipital attachment well developed, margin between it
and head concave. (12) Shaft wide proximally, distal portion of its
ventral surface concave medially. (13) Procoracoid process with apex
acute or rounded. (14) Dorsal intermuscular line swinging out to
medial surface of shaft, distal half sharply raised. (15) Ventral inter-
muscular line terminating at tip of sterno-coracoidal process. (16) Dor-
sal face moderately excavated distally, without distinct fossa. (17)
Tubercle for ligamentum sterno-coracoideum dorsale obsolete or well
developed, irregular in shape or rounded. (18) Internal distal angle


Vol. 6






HOLMAN: OSTEOLOGY OF QUAILS


without distinct tubercle or ridge on dorsal face. (19) Sterno-cora-
coidal process with distal margin slightly oblique, not produced as
sharply raised ridge, without dorsal groove; its tip directed laterally
and ending in a terminal knob. (20) Distal border of sternal facet
usually deeply concave.
SCAPULA. (21) Glenoid facet oval in dorsal view. (22) Dorsal de-
pression just media to glenoid facet moderately to deeply excavated.
(23) Bridge between acromion process and glenoid facet from less
than one-third to about one-half width of glenoid facet. (24) Acromion
process with apex rounded, usually moderately deflected.
HUMERUS. (25) Pneumatic fossa short to long, elliptical, oval, or
ovaloid, without inner shelf. (26) Median crest moderately to strongly
developed, elevated only at middle. (27) Capital groove usually mod-
erately excavated, its internal margin usually moderately elevated.
(28) Margin between head and internal tuberosity moderately concave.
(29) Head broader than long in anconal view. (30) Ridge along medial
border of fossa II slightly to moderately swollen, elongate, its apex
acute. (31) Fossa II well developed. (32) Latissimus ridge originating
on lateral edge of shaft, but swinging in onto anconal face proximally.
(33) External tuberosity moderately inflated. (34) Bicipital crest aris-
ing gently from shaft, usually narrowly rounded, its dorsal border lack-
ing a groove. (35) Deltoid crest moderately developed, slightly in-
flected, its summit flattened or knoblike. (36) Entepicondyle usually
at level of internal condyle. (37) Entepicondylar prominence moder-
ately produced, its pit on internal face. (38) Depression of M. brachi-
alis anticus usually moderately excavated. (39) Distal border of ex-
ternal condyle rounded in anconal view.
ULNA. (40) Olecranon process strongly produced, its apex pointed.
(41) External cotyla moderately elongate, rounded throughout, its dis-
tal border sloping gently into shaft, only slightly notched by proximal
radial impression. (42) Impression for M. brachialis anticus exca-
vated, its borders distinct. (43) Height at middle of shaft 62 to 71
percent of height just distal to external cotyla. (44) Distal radial im-
pression obsolete. (45) Notch between carpal tuberosity and internal
condyle well developed. (46) External condyle well produced, arising
abruptly from shaft, rounded throughout.
CARPOMETACARPUS. (47) Metacarpal I moderately pointed or
rounded terminally. (48) Ridge separating anterior carpal fossa from
excavated area above pisiform process produced, extending well onto
base of metacarpal I. (49) Pisiform process round, ovaloid, oval, or
elliptical in internal view, slightly reflected dorsad, thus usually be-
low or occasionally at level of dorsal rim of inner carpal trochlea.






BULLETIN FLORIDA STATE MUSEUM


(50) Ligamental attachment of pisiform process produced and much
swollen proximally, flattened distally, with proximal portion usually
connected to pisiform process by bony bridge. (51) Internal carpal
trochlea with proximal border shallowly concave or slightly notched,
its ventral border without notch. (52) Intermetacarpal tubercle ex-
tending to or slightly beyond, but usually not ankylosing with meta-
carpal III, distally placed with a small space between its posterior
border and junction of metacarpals II and III. (53) Tubercle on in-
ternal proximal face of metacarpal III obsolete. (54) One strongly
developed tubercle on external proximal face of metacarpal III. (55)
Metacarpal III slightly bowed.
PELVIS. (56) Width of pelvis through antitrochanters 44 to 51 per-
cent of length of innominate from anterior edge of ilium to ischial
angle. (57) Pectineal process obsolete, only moderately produced be-
yond acetabular border, its apex usually pointed. (58) Depression on
medial side of pectineal process shallowly excavated, without fora-
men. (59) Posterior iliac crest without prominence. (60) Ridge origi-
nating on posterior iliac crest at level of ilio-ischiac fenestra, continu-
ing ventrad and posteriad toward posterior border of ilium with prom-
inence sharply arising from its posterior portion. (61) Dorsal face of
postacetabular ilium moderately broad anteriad, narrowing abruptly
posteriad to form acute apex of short, broad, dorsal roof of posterior
process. (62) Caudal face of postacetabular ilium enlarged, triangular,
vertical in position, its posterodorsal apex forming posteromedial wall
of posterior process. (63) Lateral face of postacetabular ilium with
posterodorsal apex acute, forming shortened lateral wall of posterior
process. (64) Renal bar slender, both bar and posterior process exca-
vated by renal depression. (65) Ischium at angle of about 160 degrees
to ventral edge of posterior iliac crest. (66),Width through transverse
processes of fourth synsacral vertebra 57 to 68 percent of length of
synsacrum from centrum of first synsacral vertebra through transverse
process of fourth synsacral vertebra.
FEMUR. (67) Distal width 15 to 18 percent of length. (68) Head
moderately enlarged, neither bent dorsally nor rotated posteriorly.
(69) Neck only moderately long. (70) Iliac facet without compression,
posterior border convex. (71) Dorsal crest of trochanter depressed
and deflected or compressed and reflected, its dorsal border flat or
rounded. (72) Trochanteric ridge 21 to 25 percent length of femur.
(73) Distal depth 78 to 90 percent of distal width. (74) Internal con-
dyle moderately wide, its anterior border much produced throughout.
(75) External condyle compressed, rounded anteriorly.
TIBIOTARSUS. (76) Depth of inner cnemial crest 54 to 69 percent


Vol. 6






HOLMAN: OSTEOLOGY OF QUAILS


of its length. (77) Outer cnemial crest usually straight. (78) Inter-
articular tubercle usually moderately produced. (79) Area between
external articular surface and outer cnemial crest moderately concave
in dorsal view. (80) Posterior margin between external and internal
articular surface with distinct notch. (81) Area just anterior to ridge
from proximal internal ligamental attachment without deeply exca-
vated depression. (82) Internal ligamental prominence on internal
condyle moderately produced. (83) Face of internal condyle mod-
erately excavated, its border moderately elevated. (84) Dorsal border
of supratendinal bridge oblique or straight.
TARSOMETATARSUS. (85) Distal width 16 to 19 percent of length.
(86) Proximal depth 93 to 100 percent of proximal width. (87) Inter-
cotylar prominence moderately deflected, its apex usually pointed.
(88) Posterior intercotylar area weakly excavated as a triangular de-
pression. (89) Area surrounding posterior opening of inner proximal
foramen shallowly excavated. (90) Hypotarsus with 2 well-developed
and 1 obsolete calcaneal ridges, 2 calcaneal grooves, and 2 closed cal-
caneal canals. (91) Anterior face of shaft usually moderately exca-
vated by anterior metatarsal groove. (92) Depth of middle trochlea
52 to 60 percent of distal width. (93) Trochlea for digit II only slightly
inflected. (94) Wing of trochlea for digit IV depressed dorsoventrally,
thus entire trochlea much deeper than long.
INTERMEMBRAL PROPORTIONS. (95) Wing/leg 63 to 69 percent.
(96) Ulna/humerus 86 to 91 percent. (97) Tarsometatarsus/femur 73
to 83 percent. (98) Tarsometatarsus/tibiotarsus 57 to 61 percent. (99)
Humerus/femur 82 to 88 percent. (100) Humerus/tibiotarsus 62 to
66 percent. (101) Ulna/femur 72 to 79 percent. (102) Ulna/tibiotarsus
53 to 59 percent. (103) Ulna/tarsometatarsus 92 to 104 percent. (104)
Carpometacarpus/femur 43 to 47 percent. (105) Carpometacarpus/
tibiotarsus 33 to 35 percent. (106) Carpometacarpus/tarsometatarsus
55 to 62 percent. (107) Tarsometatarsus/humerus 86 to 98 percent.
(108) Ilium/sternum 66 to 75 percent. (109) Height of sternal carina/
ilium 37 to 45 percent.

SPECIFIC VARIATION IN Colinus
In this section when two of the three species agree in a particular
character, the usual condition is mentioned first, followed by the ex-
ception shown in the third species.
CORACOID. (8) Head usually moderately reflected (moderately in-
flected in C. nigrogularis). (10) Furcular facet moderately inflated
(weakly inflated in C. nigrogularis). (13) Procoracoid process with apex
usually rounded (acute in C. nigrogularis). (17) Tubercle for ligamen-






BULLETIN FLORIDA STATE MUSEUM


tum sterno-coracoideum dorsale usually obsolete and irregular in
shape (well developed and round in C. leucopogon).
SCAPULA. (23) Bridge between acromion process and glenoid facet
usually about one-half width of glenoid facet (less than one-third width
in C. nigrogularis).
HUMERUS. (25) Pneumatic fossa usually moderately long, occa-
sionally short (very long in C. nigrogularis). (26) Median crest mod-
erately developed (usually strongly developed in C. virginianus). (30)
Ridge along medial border of fossa II moderately swollen (slightly
swollen in C. nigrogularis). (35) Deltoid crest with apex flattened
(usually knoblike in C. virginianus).
CARPOMETACARPUS. (47) Metacarpal I usually moderately pointed
terminally (rounded terminally in C. leucopogon).
FEMUR. (71) Dorsal crest of trochanter usually depressed and de-
flected, its dorsal border usually flat (usually compressed and reflected,
dorsal border usually round in C. leucopogon).
TIBIOTARSUS. (84) Dorsal border of supratendinal bridge usually
oblique (straight in C. leucopogon).
INTERMEMBRAL PROPORTIONS. The species of Colinus are not sep-
arable on intermembral proportions.
REMARKS. The skeletons of Recent species of Colinus are similar.
Colinus virginianus resembles C. leucopogon in 6 of 12 characters,
C. nigrogularis in 4 characters, and is unique in 2 characters (these
are subject to some individual variation). Colinus nigrogularis resem-
bles C. virginianus in 4 of 12 characters, C. leucopogon in 2 char-
acters, and is unique in 6 characters. C. leucopogon resembles C. vir-
ginianus in 6 of 12 characters, C. nigrogularis in 2 characters and is
unique in 4 characters.
Colinus virginianus averages largest, C. leucopogon smaller, and
C. nigrogularis smallest (table 1).
SUBSPECIFIC VARIATION IN Colinus virginianus
United States populations of Recent Colinus virginianus show a grad-
ual decrease in mean skeletal size from north to south (table 2).
No qualitative skeletal differences separate the subspecies of
Colinus virginianus, although one qualitative character appears to fol-
low a north-south gradient. The Mexican samples are too small to
indicate trends.
A north-south decrease in mean size occurs in United States bob-
whites. The Michigan individuals of C. v. virginianus are largest, and
the Illinois individuals second largest in each of 20 measurements.
Northern Florida bobwhites, representing intergrades between C. v.


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HOLMAN: OSTEOLOGY OF QUAILS


virginianus and C. v. floridanus, are third largest in 19 of 20 measure-
ments, whereas the peninsular subspecies C. v. floridanus is the small-
est in 19 of 20 measurements.
The available sample of the Mexican races of C. virginianus is too
small to show size ranges adequately, but both C. v. insignis from Chi-
apas and C. v. coyolcos from Oaxaca are smaller than C. v. virginianus
from Michigan and Illinois in all 20 measurements taken.
The development of the median crest of the humerus appears to
show a north-south gradient in the subspecies of C. virginianus and
thus is the only qualitative character that may be associated with a
definite trend. In C. v. virginianus from Michigan, 85 percent of 20
specimens have the median crest strongly developed; 15 percent have
it moderately or weakly developed. In C. v. virginianus from south-
ern Illinois, 71 percent of 38 specimens have the median crest strongly
developed; 29 percent have it moderately or weakly developed. In
intergrades between C. v. virginianus and C. v. floridanus from north-
ern Florida, 38 percent of 13 specimens have the median crest strong-
ly developed; in 62 percent it is moderately or weakly developed. In
C. v. floridanus from peninsular Florida, 29 percent of 7 specimens
have the median crest strongly developed; 71 percent have it mod-
erately or weakly developed. In C. v. insignis from Chiapas, Mexico,
40 percent of 5 specimens have the median crest strongly developed;
60 percent are moderately developed. One specimen of C. v. coyolcos
from Oaxaca, Mexico, has the median crest strongly developed.

Lophortyx BONAPARTE, 1838
The genus Lophortyx occurs in the western United States and western
Mexico. Table 1 includes measurements of the three recognized spe-
cies, L. californica (Shaw, 1798), L. gambelii Gambel, 1843 and L.
douglasii (Vigors, 1829).
RosTRUM. (1) Relatively long and shallow. (2) Nasal fossae el-
liptical.
STERNUM. (3) Height of carina through sternal plate 28 to 31
percent length of sternum. (4) Dorsal lip of coracoidal sulcus projected
well beyond ventral lip. (5) Depressions on anterior sternal plate ob-
solete or deep, slightly to moderately pneumatic. (6) Base of posterior
lateral process less than one-half width of sternal plate at its middle.
CORACOm. (7) Entire proximal end straight above procoracoid
process. (8) Head narrowly rounded, moderately compressed, mod-
erately inflected. (9) Brachial tuberosity with slight dorsal and ven-
tral overhang. (10) Furcular facet weakly inflated. (11) Bicipital






BULLETIN FLORIDA STATE MUSEUM


attachment well developed, margin between it and head concave.
(12) Shaft narrow or wide proximally, distal portion of ventral sur-
face convex or concave medially. (13) Procoracoid process with apex
rounded. (14) Dorsal intermuscular line swinging out to medial sur-
face of shaft, distal half sharply raised. (15) Ventral intermuscular line
terminating at tip of sterno-coracoidal process. (16) Dorsal face
weakly or moderately excavated distally, without fossa. (17) Tubercle
for ligamentum sterno-coracoideum dorsale obsolete or well devel-
oped, round. (18) Internal distal angle without distinct tubercle or
ridge on dorsal face. (19) Sterno-coracoidal process with distal margin
slightly oblique, not produced as sharply raised ridge, without dorsal
groove; its tip directed laterally and ending in terminal knob. (20)
Distal border of sternal facet deeply to very deeply concave.
SCAPULA. (21) Glenoid facet oval in dorsal view. (22) Dorsal de-
pression just media to glenoid facet shallowly to moderately exca-
vated. (23) Bridge between acromion process and glenoid facet about
one-half or more width of glenoid facet. (24) Acromion process with
apex rounded and usually slightly deflected.
HUMERUS. (25) Pneumatic fossa a short to moderately long oval
or ellipse, without inner shelf. (26) Median crest weak to strongly
developed, elevated throughout or only at its middle. (27) Capital
groove moderately excavated, its internal margin moderately elevated.
(28) Margin between head and internal tuberosity moderately con-
cave. (29) Head as broad or broader than long in anconal view. (30)
Ridge along medial border of fossa II moderately swollen, elongate,
its apex acute. (31) Fossa II well developed. (32) Latissimus ridge
originating on lateral edge of shaft, but swinging in onto anconal
face proximally. (33) External tuberosity moderately inflated. (34)
Bicipital crest arising gently from shaft, narrowly rounded, its dis-
tal border lacking a groove. (35) Deltoid crest moderately devel-
oped, slightly inflected, its summit usually flattened. (36) Entepi-
condyle produced slightly beyond internal condyle. (37) Entepicondy-
lar prominence moderately produced, its pit on internal face. (38) De-
pression of M. brachialis anticus moderately excavated. (39) Distal
border of external condyle rounded in anconal view.
ULNA. (40) Olecranon process strongly produced, its apex pointed.
(41) External cotyla moderately elongate, rounded throughout, its dis-
tal border sloping gently into shaft, usually only slightly notched by
proximal radial impression. (42) Impression for M. brachialis anticus
excavated, its borders distinct. (43) Height at middle of shaft 61 to
72 percent of height just distal to external cotyla. (44) Distal radial
impression obsolete. (45) Notch between carpal tuberosity and in-


Vol. 6






HOLMAN: OSTEOLOGY OF QUAILS


ternal condyle well developed. (46) External condyle well produced,
arising gently from shaft, rounded throughout.
CARPOMETACARPUS. (47) Metacarpal I moderately pointed or
rounded terminally. (48) Ridge separating anterior carpal fossa from
excavated area above pisiform process produced, extending well onto
base of metacarpal I. (49) Pisiform process ovaloid in internal view,
slightly or moderately reflected dorsad, thus below, at, or slightly
above level of dorsal rim of inner carpal trochlea. (50) Ligamental
attachment of pisiform process produced and much swollen proximal-
ly, flattened distally, with proximal portion usually connected to pisi-
form process by bony bridge. (51) Internal carpal trochlea with prox-
imal border usually slightly notched, its ventral border without notch.
(52) Intermetacarpal tubercle extending to and sometimes ankylosing
with metacarpal III, distally placed with a small space between its
posterior border and junction of metacarpals II and III. (53) Tubercle
on internal proximal face of metacarpal III obsolete. (54) One strong-
ly developed tubercle on external proximal face of metacarpal III.
(55) Metacarpal III slightly bowed.
PELVIs. (56) Width of pelvis through antitrochanters 54 to 57
percent of length of innominate from anterior edge of ilium to ischial
angle. (57) Pectineal process obsolete, only moderately produced be-
yond acetabular border, its apex pointed. (58) Depression on medial
side of pectineal process shallowly excavated, without foramen. (59)
Posterior iliac crest without prominence. (60) Ridge originating on
posterior iliac crest at level of ilio-ischiac fenestra, continuing ven-
trad and posteriad toward posterior border of ilium with prominence
sharply arising from posterior portion. (61) Dorsal face of postace-
tabular ilium moderately broad anteriad, narrowing abruptly posteriad
to form acute apex of short, broad, dorsal roof of posterior process.
(62) Caudal face of postacetabular enlarged, triangular, vertical in
position, its posterodorsal apex forming posteromedial wall of posterior
process. (63) Lateral face of postacetabular ilium with posterodorsal
apex acute, forming shortened lateral wall of posterior process. (64)
Renal bar slender, with both bar and posterior process excavated by
renal depression. (65) Ischium at angle of about 160 degrees to ven-
tral edge of posterior iliac crest. (66) Width through transverse pro-
cess of fourth synsacral vertebra 67 to 76 percent of length of synsac-
rum from centrum of first synsacral vertebra through transverse process
of fourth synsacral vertebra.
FEMUR. (67) Distal width 17 to 18 percent of length. (68) Head
moderately or much enlarged, neither bent dorsally nor rotated pos-
teriorly. (69) Neck only moderately long. (70) Iliac facet without







BULLETIN FLORIDA STATE MUSEUM


compression, posterior border convex. (71) Dorsal crest of trochanter
usually compressed and reflected, its dorsal border rounded. (72) Tro-
chanteric ridge 24 to 25 percent length of femur. (73) Distal depth
77 to 85 percent of distal width. (74) Internal condyle moderately
wide, its anterior border much produced throughout. (75) External
condyle compressed, rounded anteriorly.
TIBIOTARSUS. (76) Depth of inner cnemial crest 58 to 67 percent
of its length. (77) Outer cnemial crest straight or decurved. (78) In-
terarticular tubercle moderately produced. (79) Area between external
articular surface and outer cnemial crest moderately concave in dor-
sal view. (80) Posterior margin between external and internal articu-
lar surfaces with distinct notch. (81) Area just anterior to ridge from
proximal internal ligamental attachment without deeply excavated
depression. (82) Internal ligamental prominence on internal condyle
moderately produced. (83) Face of internal condyle moderately exca-
vated, its border moderately elevated. (84) Dorsal border of supra-
tendinal bridge usually straight.
TARSOMETATARSUS. (85) Distal width 18 to 20 percent of length.
(86) Proximal depth 93 to 100 percent of proximal width. (87) Inter-
cotylar prominence moderately deflected, its apex pointed. (88) Pos-
terior intercotylar area usually weakly excavated as a triangular de-
pression. (89) Area surrounding posterior opening of inner proximal
foramen shallowly excavated. (90) Hypotarsus with 2 well-developed
and 1 obsolete calcaneal ridges, 2 calcaneal grooves, and 2 closed cal-
caneal canals. (91) Anterior metatarsal groove usually reduced. (92)
Depth of middle trochlea about 51 to 57 percent of distal width. (93)
Trochlea for digit II only slightly inflected. (94) Wing of trochlea
for digit IV depressed dorsoventrally, thus entire trochlea much deeper
than long.
INTERMEMBRAL PROPORTIONS. (95) Wing/leg 60 to 66 percent.
(96) Ulna/humerus 86 to 89 percent. (97) Tarsometatarsus/femur 76
to 82 percent. (98) Tarsometatarsus/tibiotarsus 57 to 59 percent. (99)
Humerus/femur 79 to 85 percent. (100) Humerus/tibiotarsus 58 to
65 percent. (101) Ulna/femur 68 to 74 percent. (102) Ulna/tibiotar-
sus 50 to 56 percent. (103) Ulna/tarsometatarsus 87 to 97 percent.
(104) Carpometacarpus/femur 37 to 45 percent. (105) Carpometacar-
pus/tibiotarsus 31 to 35 percent. (106) Carpometacarpus/tarsometa-
tarsus 54 to 60 percent. (107) Tarsometatarsus/humerus 89 to 99 per-
cent. (108) Ilium/sternum 66 to 70 percent. (109) Height of sternal
carina/ilium 42 to 45 percent.


Vol. 6







HOLMAN: OSTEOLOGY OF QUAILS


SPECIFIC VARIATION IN Lophortyx
In this section when two of the three species agree in a particular
character, this condition is mentioned first, followed in parentheses
by the condition shown by the third species.
STERNUM. (5) Depressions on anterior portion of sternal plate
deep (obsolete in L. californica); slightly to moderately pneumatic in
L. douglasii, slightly pneumatic in L. californica, moderately pneumatic
in L. gambelii.
CORACOID. (12) Shaft wide proximally (narrow proximally in L.
californica) distal portion of ventral surface concave medially (convex
medially in L. gambelii). (16) Dorsal face weakly excavated distally
(moderately excavated distally in L. douglasii). (17) Tubercle for lig-
amentum stero-coracoideum dorsale well developed (obsolete in L.
californica). (20) Distal border of sternal facet deeply concave (very
deeply concave in L. gambelii).
SCAPULA. (22) Dorsal depression just media to glenoid facet
shallowly to moderately excavated in L. douglasii, shallowly excavated
in L. californica, moderately excavated in L. gambelii. (23) Bridge
between acromion process and glenoid facet about one-half width of
glenoid facet (more than one-half width of glenoid facet in L. gam-
belii).
HUMERUS. Combined proximal width, middle shaft width, and
distal width, 58 to 59 percent of length (60 to 63 percent of length in
L. californica). Proximal width 27 percent of length (28 to 30 percent
of length in L. californica). (26) Median crest strongly developed, ele-
vated only at its middle (weak and elevated throughout in L. douglasii).
(29) Head broader than long in anconal view (as broad as long in L.
gambelii).
ULNA. (43) Height at middle of shaft 67 to 72 percent of height
just distal to external cotyla (61 percent of height in L. gambelii).
CARPOMETACARPUS. Proximal width through pisiform process 65
to 67 percent of proximal height through metacarpal I (61 to 62 per-
cent of proximal height in L. californica). (47) Metacarpal I rounded
terminally (moderately pointed in L. californica). (49) Pisiform pro-
cess slightly reflected dorsad, thus below level of dorsal rim of inner
carpal trochlea (moderately reflected dorsad, thus at or slightly above
dorsal rim of inner carpal trochlea in L. gambelii). (52) Intermeta-
carpal tubercle usually extending to and ankylosing with metacarpal
III (extending to, but not ankylosing with metacarpal III in L. doug-
lasii).
FEMUR. (68) Head moderately large (much enlarged in L. gam-
belii).







BULLETIN FLORIDA STATE MUSEUM


TIBIOTARSUS. Proximal depth 76 to 77 percent of proximal width
(72 to 73 percent of proximal width in L. californica). (77) Outer
cnemial crest straight or slightly decurved in L. californica, straight
in L. douglasii, decurved in L. gambelii.
INTERMEMBRAL PROPORTIONS. (95) Wing/leg 60 to 63 percent (66
percent in L. douglasii). Femur/tibiotarsus 72 to 74 percent (76 to 77
percent in L. douglasii). (97) Tarsometatarsus/femur 76 to 79 percent
(82 percent in L. gambelii). (99) Humerus/femur 83 to 85 percent
(79 to 80 percent in L. californica). (100) Humerus/tibiotarsus 58 to 60
percent (64 to 65 percent in L. douglasii). (102) Ulna/tibiotarsus 50
to 53 percent (55 to 56 percent in L. douglasii). (103) Ulna/tarso-
metatarsus 87 to 90 percent (97 percent in L. douglasii). (104) Carpo-
metacarpus/femur 45 percent (37 to 42 percent in L. californica).
(105) Carpometacarpus/tibiotarsus 31 to 32 percent (34 to 35 percent
in L. douglasii). (106) Carpometacarpus/tarsometatarsus 54 to 56
percent (59 to 60 percent in L. douglasii). (107) Tarsometatarsus/
humerus 98 to 99 percent (89 to 90 percent in L. douglasii).

REMARKS. On the basis of the samples studied, the skeletons of Re-
cent species of Lophortyx show more interspecific differences than do
those of Recent species of Colinus. Lophortyx californica resembles
L. gambelii in 11 of 30 characters, L. douglasii in 8 characters, and
is unique in 11 characters. Lophortyx gambelii resembles L. cali-
fornica in 11 of 30 characters, L. douglasii in 10 characters, and is
unique in 9 characters. L. douglasii resembles L. californica in 8 of 30
characters, L. gambelii in 10 characters, and is unique in 12 characters.
Lophortyx californica and L. gambelii are similar in size, but L.
douglasii averages somewhat smaller in most measurements (table 1).

Odontophorus VIEILLOT, 1816
The genus Odontophorus is confined to tropical America. Of the 16
species recognized, 0. gujanensis (Gmelin, 1789), 0. stellatus (Gould,
1843), and 0. guttatus (Gould, 1838) were examined;, their measure-
ments are given in table 1.
ROSTRUM. (1) Relatively long and shallow. (2) Nasal fossae el-
liptical.
STERNUM. (3) Height of carina through sternal plate 29 to 33
percent length of sternum. (4) Dorsal lip of coracoidal sulcus projected
slightly or well beyond ventral lip. (5) Depressions on anterior por-
tion of sternal plate obsolete and nonpneumatic. (6) Base of posterior
lateral process less than one-half width of sternal plate at middle.


Vol. 6






HOLMAN: OSTEOLOGY OF QUAILS


CORACOID. (7) Entire proximal end straight above procoracoid
process. (8) Head narrowly rounded, moderately compressed, mod-
erately reflected or inflected. (9) Brachial tuberosity with slight or
moderate dorsal, and slight ventral overhang. (10) Furcular facet
much inflated. (11) Bicipital attachment well developed, margin be-
tween it and head concave. (12) Shaft wide proximally, distal portion
of ventral surface concave medially. (13) Procoracoid process with
apex rounded. (14) Dorsal intermuscular line swinging out to medial
surface of shaft, distal two-thirds or three-fourths sharply raised. (15)
Ventral intermuscular line terminating at tip of sterno-coracoidal proc-
ess. (16) Dorsal face moderately or deeply excavated distally, with-
out fossa. (17) Tubercle for ligamentum sterno-coracoideum dorsale
obsolete and irregular in shape. (18) Internal distal angle without
distinct tubercle or ridge on dorsal face. (19) Sterno-coracoidal process
with distal margin slightly oblique, not produced as sharply raised
ridge, without dorsal groove; its tip directed laterally and ending in
a terminal knob. (20) Distal border of sternal facet shallowly or deeply
concave.
SCAPULA. (21) Glenoid facet oval or elliptical in dorsal view. (22)
Dorsal depression just media to glenoid facet shallowly to deeply
excavated. (23) Bridge between acromion process and glenoid facet
usually about one-half width of glenoid facet. (24) Acromion process
with apex rounded, slightly or moderately deflected.
HUMERUS. (25) Pneumatic fossa short to moderately long, oval
or elliptical, without inner shelf, or with perforated inner shelf ex-
tending from external bicipital surface to medial bar. (26) Median
crest strongly or very strongly developed, elevated throughout. (27)
Capital groove moderately excavated, its internal margin moderately
elevated. (28) Margin between head and internal tuberosity moder-
ately concave. (29) Head broader than long in anconal view. (30)
Ridge along medial border of fossa II moderately swollen, elongate,
its apex acute. (31) Fossa II obsolete. (32) Latissimus ridge originat-
ing on lateral edge of shaft, but swinging in onto anconal face prox-
imally. (33) External tuberosity slightly, moderately, or much inflated.
(34) Bicipital crest arising gently from shaft, rounded, its distal border
lacking a groove. (35) Deltoid crest moderately or strongly developed,
slightly inflected, its summit knoblike. (36) Entepicondyle produced
slightly beyond level of internal condyle. (37) Entepicondylar prom-
inence much produced, its pit on internal face. (38) Depression of M.
brachialis anticus moderately excavated. (39) Distal border of ex-
ternal condyle rounded in anconal view.





BULLETIN FLORIDA STATE MUSEUM


ULNA. (40) Olecranon process strongly produced, its apex pointed.
(41) External cotyla moderately elongate, rounded throughout or flat-
tened dorsally, its distal border sloping gently or precipitously into
shaft, slightly or deeply notched by proximal radial impression. (42)
Impression for M. brachialis anticus excavated, its borders distinct.
(43) Height at middle of shaft 65 to 72 percent of height just distal to
external cotyla. (44) Distal radial impression obsolete or moderately
excavated. (45) Notch between carpal tuberosity and internal con-
dyle well developed. (46) External condyle well produced, arising
abruptly from shaft, rounded throughout.
CARPOMETACARPUS. (47) Metacarpal I moderately pointed termi-
nally. (48) Ridge separating anterior carpal fossa from excavated area
above pisiform process produced, extending well onto base of meta-
carpal I. (49) Pisiform process ovaloid in internal view, slightly re-
flected dorsad, thus slightly below level of dorsal rim of inner carpal
trochlea. (50) Ligamental attachment of pisiform process produced,
moderately or much swollen proximally, flattened distally, with prox-
imal portion either connected to pisiform process by bony bridge or
separated from pisiform process by shallow sulcus. (51) Internal car-
pal trochlea with proximal border usually slightly notched, its ventral
border without notch. (52) Intermetacarpal tubercle extending to but
not ankylosing with metacarpal III, distally placed with a small space
between its posterior border and junction of metacarpals II and III.
(53) Tubercle on internal proximal face of metacarpal III obsolete.
(54) One strongly developed tubercle on external proximal face of
metacarpal III. (55) Metacarpal III slightly bowed.
PELVIS. (56) Width of pelvis through antitrochanters 51 to 53
percent of length of innominate from anterior edge of ilium to ischial
angle. (57) Pectineal process obsolete, only slightly produced beyond
acetabular border as a minute point. (58) Depression on medial side
of pectineal process obsolete, without foramen. (59) Posterior iliac
crest usually without prominence. (60) Ridge originating on posterior
iliac crest at level of ilio-ischiac fenestra, continuing ventrad and pos-
teriad toward posterior border of ilium with prominence weakly or
sharply arising from posterior portion; or pelvis without this ridge or
prominence. (61) Dorsal face of postacetabular ilium moderately
broad anteriad, narrowing posteriad to form rounded or pointed apex
of moderately or much elongated, narrow, dorsal roof of posterior
process. (62) Caudal face of postacetabular ilium obsolete, triangular,
horizontal in position, forming ventral floor of posterior process. (63)
Lateral face of postacetabular ilium with posterodorsal apex acute,
forming moderately or much elongated lateral wall of posterior proc-


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HOLMAN: OSTEOLOGY OF QUAILS


ess. (64) Renal bar broad, bar, but not posterior process, excavated by
renal depression. (65) Ischium at angle of about 160 degrees to ven-
tral edge of posterior iliac crest. (66) Width through transverse proc-
esses of fourth synsacral vertebra 71 to 79 percent length of synsacrum
from centrum of first synsacral vertebra through transverse process
of fourth synsacral vertebra.
FEMUR. (67) Distal width 17 to 19 percent of length. (68) Head
moderately enlarged, neither bent dorsally nor rotated posteriorly.
(69) Neck only moderately long. (70) Iliac facet without compression,
posterior border convex. (71) Dorsal crest of trochanter depressed
and deflected, its dorsal border flat. (72) Trochanteric ridge 21 to 26
percent length of femur. (73) Distal depth 83 to 85 percent of distal
width. (74) Internal condyle moderately wide, its anterior border
much produced throughout. (75) External condyle compressed, round-
ed anteriorly.
TIBIOTARSUS. (76) Depth of inner cnemial crest 58 to 68 percent
of its length. (77) Outer cnemial crest straight. (78) Interarticular
tubercle moderately or strongly produced. (79) Area between ex-
ternal articular surface and outer cnemial crest moderately or deeply
concave in dorsal view. (80) Posterior margin between external and
internal articular surface with distinct notch. (81) Area just anterior
to ridge for proximal internal ligamental attachment without deeply
excavated depression. (82) Internal ligamental prominence on inter-
nal condyle moderately produced. (83) Face of internal condyle mod-
erately excavated, its border moderately elevated. (84) Dorsal border
of supratendinal bridge usually straight.
TARSOMETATARSUS. (85) Distal width 17 to 18 percent of length.
(86) Proximal depth 96 to 100 percent of proximal width. (87) Inter-
cotylar prominence moderately deflected, its apex pointed. (88) Pos-
terior intercotylar area moderately excavated as triangular depression.
(89) Area surrounding posterior opening of inner proximal foramen
usually deeply excavated. (90) Hypotarsus with 2 well-developed and
1 obsolete calcaneal ridges, 2 calcaneal grooves, and 1 or 2 closed
calcaneal canals. (91) Anterior metatarsal groove reduced. (92)
Depth of middle trochlea 54 to 57 percent of distal width. (93) Tro-
chlea for digit II only slightly inflected. (94) Wing of trochlea for
digit IV depressed anteroposteriorly, thus entire trochlea slightly long-
er than deep.
INTERMEMBRAL PROPORTIONS. (95) Wing/leg 64 to 68 percent.
(96) Ulna/humerus 96 to 102 percent. (97) Tarsometatarsus/femur 84
to 87 percent. (98) Tarsometatarsus/tibiotarsus 63 to 65 percent. (99)
Humerus/femur 82 to 86 percent. (100) Humerus/tibiotarsus 61 to





BULLETIN FLORIDA STATE MUSEUM


64 percent. (101) Ulna/femur 80 to 85 percent. (102) Ulna/tibiotarsus
60 to 62 percent. (103) Ulna/tarsometatarsus 92 to 99 percent. (104)
Carpometacarpus/femur 45 to 49 percent. (105) Carpometacarpus/
tibiotarsus 33 to 36 percent. (106) Carpometacarpus/tarsometatarsus
51 to 58 percent. (107) Tarsometatarsus/humerus 99 to 101 percent.
(108) Ilium/sternum 71 to 76 percent. (109) Height of sternal carina/
ilium 41 to 44 percent.
SPECIFIC VARIATION IN Odontophorus
When two of three species agree in a particular character, this condi-
tion is mentioned first, followed by the condition shown by the third
species.
STERNUM. (3) Height of carina through sternal plate 32 to 33 per-
cent length of sternum (29 percent length of sternum in 0. guttatus).
(4) Dorsal lip of coracoidal sulcus projected well beyond ventral lip
(projected only slightly beyond ventral lip in 0. gujanensis).
CORACOID. (8) Head moderately reflected (moderately inflected
in 0. guttatus). (9) Brachial tuberosity with moderate dorsal over-
hang (slight dorsal overhang in 0. gujanensis). (14) Dorsal intermus-
cular line with distal three-fourths sharply raised (with distal two-
thirds sharply raised in 0. guttatus). (16) Dorsal face deeply exca-
vated distally (moderately excavated distally in 0. stellatus). (20)
Distal border of sternal facet shallowly concave (deeply concave in
0. guttatus).
SCAPULA. (21) Glenoid facet oval in dorsal view (elliptical in O.
guttatus). (22) Dorsal depression just media to glenoid facet mod-
erately or deeply excavated shallowlyy excavated in 0. stellatus). (24)
Acromion process with apex only slightly deflected (moderately de-
flected in 0. guttatus).
HUMERUS. (25) Pneumatic fossa oval, without inner shelf (ellipti-
cal, with perforated inner shelf extending from external bicipital sur-
face to medial bar in 0. gujanensis). (26) Median crest strongly devel-
oped (very strongly developed in 0. gujanensis). (33) External tuber-
osity slightly inflated in 0. guttatus, moderately inflated in 0. stellatus,
much inflated in 0. gujanensis. (35) Deltoid crest moderately devel-
oped (strongly developed in 0. gujanensis).
ULNA. (41) External cotyla rounded. throughout (flattened dor-
sally in 0. guttatus), its distal border sloping gently into shaft (sloping
precipitously in 0. guttatus), only slightly notched by proximal radial
impression (deeply notched in 0. gujanensis). (44) Distal radial im-
pression obsolete (moderately excavated in 0. gujanensis).
CARPOMETACARPUS. (50) Ligamental attachment of pisiform proc-


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HOLMAN: OSTEOLOGY OF QUAILS


ess much swollen proximally (moderately swollen proximally in O.
gujanensis).
PELVIS. (60) Ridge originating on posterior iliac crest at level
of ilio-ischiac fenestra, continuing ventrad and posteriad toward pos-
terior border of ilium with prominence sharply arising from its poste-
rior portion in 0. guttatus (prominence weakly arising in 0. stellatus,
sharply arising, or without ridge and prominence in 0. gujanensis).
(61) Dorsal face of postacetabular ilium much elongated and narrow,
its apex pointed (moderately elongated and narrow, its apex rounded
in 0. stellatus). (63) Lateral face of postacetabular ilium much elon-
gated (moderately elongate in 0. guttatus).
TIBIOTARSUS. (76) Depth of inner cnemial crest 65 to 68 percent
of its length (58 percent of its length in 0. gujanensis). (78) Interartic-
ular tubercle moderately produced (strongly produced in 0. stellatus).
(79) Area between external articular surface and outer cnemial crest
deeply concave in dorsal view (moderately concave in 0. gujanensis).
TARSOMETATARSUS. (90) Hypotarsus with 1 closed calcaneal canal
in 0. stellatus, 1 or 2 in 0. gujanensis, 2 in 0. guttatus.
INTERMEMBRAL PROPORTIONS. (95) Wing/leg 67 to 68 percent (64
percent in 0. guttatus). Carpometacarpus/humerus 57 percent (54
percent in 0. guttatus). (98) Tarsometatarsus/tibiotarsus 63 percent
(65 percent in 0. guttatus). (99) Humerus/femur 84 to 86 percent
(82 percent in 0. guttatus). (101) Ulna/femur 83 to 85 percent (80
percent in 0. guttatus). (103) Ulna/tarsometatarsus 98 to 99 percent
(92 percent in 0. guttatus). (104) Carpometacarpus/femur 47 to 49
percent (45 percent in 0. guttatus). (105) Carpometacarpus/tibiotar-
sus 36 percent (33 percent in 0. guttatus). (106) Carpometacarpus/
tarsometatarsus 55 to 58 percent (51 percent in 0. guttatus). (108)
Ilium/sternum 74 to 76 percent (71 percent in 0. guttatus).
REMARKS. As in skeletons of Recent Lophortyx, those of Odonto-
phorus show more interspecific differences than do those of Recent
species of Colinus. Odontophorus guttatus differs from 0. gujanensis
and 0. stellatus in many intermembral ratios, and thus has more unique
characters. Odontophorus gujanensis resembles 0. stellatus in 19 of 33
characters, 0. guttatus in 3 characters, and is unique in 11 characters.
Odontophorus stellatus resembles 0. gujanensis in 19 of 33 characters,
0. guttatus in 10 characters, and is unique in 4 characters. 0. guttatus
resembles 0. stellatus in 10 of 33 characters, 0. gujanensis in 3 char-
acters, and is unique in 20 characters.
According to the small available sample, the skeletons of O. gu-
janensis and 0. guttatus are similar in size and slightly larger than O.
stellatus.





BULLETIN FLORIDA STATE MUSEUM


Dactylortyx OGILVIE-GRANT, 1893
The single species of this genus, Dactylortyx thoracicus (Gambel,
1848) is confined to Mexico and northern Central America, mainly in
the mountains. Measurements are given in table 1.
ROSTRUM. (1) Relatively long and shallow. (2) Nasal fossae el-
liptical.
STERNUM. (3) Height of carina through sternal plate 31 to 34 per-
cent length of sternum. (4) Dorsal lip of coracoidal sulcus even with
or projecting only slightly beyond ventral lip. (5) Depressions on an-
terior part of sternal plate shallow or obsolete, nonpneumatic. (6)
Base of posterior lateral process less than one-half width of sternal
plate at its middle.
CORACOID. (7) Entire proximal end straight above procoracoid
process. (8) Head narrowly rounded, moderately compressed, mod-
erately inflected. (9) Brachial tuberosity with slight dorsal and ven-
tral overhang. (10) Furcular facet moderately inflated. (11) Bicipital
attachment well developed, margin between it and head concave.
(12) Shaft wide proximally, distal portion of ventral surface concave
medially. (13) Procoracoid process with apex rounded. (14) Dorsal
intermuscular line swinging out to medial surface of shaft, distal
three-fourths sharply raised. (15) Ventral intermuscular line terminat-
ing at tip of sterno-coracoid process. (16) Dorsal face deeply exca-
vated distally, without fossa. (17) Tubercle for ligamentum sterno-
coracoideum dorsale usually obsolete, always irregular in shape. (18)
Internal distal angle without distinct tubercle or ridge. (19) Sterno-
coracoidal process with distal margin slightly oblique, not produced
as a sharply raised ridge, without dorsal groove; its tip directed lat-
erally and ending in a terminal knob. (20) Distal border of sternal
facet deeply concave.
SCAPULA. (21) Glenoid facet oval in dorsal view. (22) Dorsal de-
pression just media to glenoid facet shallowly excavated. (23) Bridge
between acromion process and glenoid facet about one-half width
of glenoid facet. (24) Acromion process with apex rounded, slight to
moderate deflection.
HUMERUS. (25) Pneumatic fossa a short to moderately long ellipse
or ovaloid, without inner shelf. (26) Median crest strongly developed,
elevated only at its middle. (27) Capital groove moderately excavated,
its internal margin moderately elevated. (28) Margin between head
and internal tuberosity moderately concave. (29) Head broader than
long in anconal view. (30) Ridge along medial border of fossa II
much swollen, shortened, its apex rounded. (31) Fossa II well de-


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HOLMAN: OSTEOLOGY OF QUAILS


veloped. (32) Latissimus ridge originating on lateral edge of shaft,
but swinging in onto anconal face proximally. (33) External tuberosity
moderately inflated. (34) Bicipital crest arising gently from shaft,
pointed, its distal border lacking a groove. (35) Deltoid crest moder-
ately developed, slightly inflected, its summit knoblike. (36) Entepi-
condyle usually at level of internal condyle. (37) Entepicondylar
prominence much produced, its pit usually on internal face. (38) De-
pression of M. brachialis anticus shallowly excavated. (39) Distal bor-
der of external condyle rounded in anconal view.
ULNA. (40) Olecranon process strongly produced, its apex trun-
cated. (41) External cotyla moderately elongate, rounded throughout,
its distal border sloping gently into shaft, only slightly notched by
proximal radial impression. (42) Impression for M. brachialis anticus
excavated, its borders distinct. (43) Height at middle of shaft 71 per-
cent of height just distal to external cotyla. (44) Distal radial impres-
sion obsolete. (45) Notch between carpal tuberosity and internal
condyle well developed. (46) External condyle moderately produced,
usually arising abruptly from shaft, usually rounded throughout.
CARPOMETACARPUS. (47) Metacarpal I usually rounded terminally.
(48) Ridge separating anterior carpal fossa from excavated area above
pisiform process produced, extending well onto base of metacarpal I.
(49) Pisiform process ovaloid in internal view, slightly reflected dorsad,
thus below level of dorsal rim of inner carpal trochlea. (50) Ligamen-
tal attachment of pisiform process produced and much swollen prox-
imally, flattened distally, with proximal portion connected to pisiform
process by bony bridge. (51) Internal carpal trochlea with proximal
border slightly notched, its ventral border without notch. (52) Inter-
metacarpal tubercle extending to but not ankylosing with metacarpal
III, proximally placed with only a minute space between its posterior
border and junction of metacarpals II and III. (53) Tubercle on in-
ternal proximal face of metacarpal III obsolete. (54) One moderately
developed tubercle on external proximal face of metacarpal III. (55)
Metacarpal III slightly bowed.
PELVIS. (56) Width of pelvis through antitrochanters 55 to 58 per-
cent length of innominate from anterior edge of ilium to ischial angle.
(57) Pectineal process absent, or obsolete and only slightly produced
beyond acetabular border as a minute point. (58) Depression on me-
dial side of pectineal process absent. (59) Posterior iliac crest with
prominence moderately produced from just posterior to ilio-ischiac
fenestra. (60) No ridge or prominence ventral to posterior iliac crest.
(61) Dorsal face of postacetabular ilium moderately broadened anteri-
ad, narrowing gently posteriad to form truncated apex of moderately





BULLETIN FLORIDA STATE MUSEUM


long, narrow, dorsal roof of posterior process. (62) Caudal face of post-
acetabular ilium obsolete, rectangular, horizontal in position, forming
ventral floor of posterior process. (63) Lateral face of postacetabular
ilium with its posterodorsal apex acute, forming elongate lateral wall
of posterior process. (64) Renal bar broad, bar, but not posterior proc-
ess, excavated by renal depression. (65) Ischium at angle of about
160 degrees to ventral edge of posterior iliac crest. (66) Width through
transverse process of fourth synsacral vertebra 88 to 93 percent of
length of synsacrum from centrum of first synsacral vertebra through
transverse process of fourth synsacral vertebra.
FEMUR. (67) Distal width 19 percent of length. (68) Head much
enlarged, neither bent dorsally nor rotated posteriorly. (69) Neck only
moderately long. (70) Iliac facet without compression, with posterior
border convex. (71) Dorsal crest of trochanter compressed and re-
flected, its dorsal border rounded. (72) Trochanteric ridge 15 to 18
percent length of femur. (73) Distal depth 81 to 84 percent of distal
width. (74) Internal condyle narrow, its anterior border usually weak-
ly produced throughout. (75) External condyle compressed, rounded
anteriorly.
TIBIOTARSUS. (76) Depth of inner cnemial crest 42 to 48 percent
of its length. (77) Outer cnemial crest straight. (78) Interarticular
tubercle weakly produced. (79) Area between external articular sur-
face and outer cnemial crest deeply concave in dorsal view. (80) Pos-
terior margin between external and internal articular surface with
distinct notch. (81) Area just anterior to ridge for proximal internal
ligamental attachment without deeply excavated depression. (82)
Area of internal ligamental attachment on internal condyle not pro-
duced as a prominence. (83) Face of internal condyle moderately ex-
cavated, its border moderately elevated. (84) Dorsal border of supra-
tendinal bridge straight.
TARSOMETATARSUS. (85) Distal width 19 to 21 percent of length.
(86) Proximal depth 88 to 94 percent of proximal width. (87) Inter-
cotylar prominence only slightly deflected, its apex usually pointed.
(88) Posterior intercotylar area moderately excavated as a triangular
depression. (89) Area surrounding posterior opening of inner proximal
foramen moderately excavated. (90) Hypotarsus with 2 well-devel-
oped and 1 weakly developed calcaneal ridges, 2 calcaneal grooves,
and 2 closed calcaneal canals. (91) Anterior metatarsal groove reduced
or obsolete. (92) Depth of middle trochlea 45 to 50 percent of distal
width. (93) Trochlea for digit II moderately inflected. (94) Wing of
trochlea for digit IV depressed dorsoventrally, thus entire trochlea
much deeper than long.


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HOLMAN: OSTEOLOGY OF QUAILS


INTERMEMBRAL PROPORTIONS. (95) Wing/leg 72 to 73 percent.
(96) Ulna/humerus 98 to 100 percent. (97) Tarsometatarsus/femur
81 to 82 percent. (98) Tarsometatarsus/tibiotarsus 59 to 60 percent.
(99) Humerus/femur 90 to 91 percent. (100) Humerus/tibiotarsus 66
to 67 percent. (101) Ulna/femur 88 to 91 percent. (102) Ulna/tibio-
tarsus 65 to 66 percent. (103) Ulna/tarsometatarsus 109 to 111 percent.
(104) Carpometacarpus/femur 48 to 50 percent. (105) Carpometacar-
pus/tibiotarsus 35 to 36 percent. (106) Carpometacarpus/tarsometa-
tarsus 59 to 62 percent. (107) Tarsometatarsus/humerus 90 percent.
(108) Ilium/sternum 70 to 78 percent. (109) Height of sternal carina/
ilium 40 to 49 percent.

Cyrtonyx GOULD, 1844
The genus Cyrtonyx is confined to the mountains of the southwestern
United States, Mexico, and northern Central America. Three species
are recognized, of which only C. montezumae (Vigors, 1830) was ex-
amined; its measurements are given in table 1.
ROSTRUM. (1) Relatively long and shallow. (2) Nasal fossae el-
liptical.
STERNUM. (3) Height of carina through sternal plate 28 to 32 per-
cent length of sternum. (4) Dorsal lip of coracoidal sulcus usually pro-
jected well beyond ventral lip. (5) Depressions on anterior part of
sternal plate shallow and nonpneumatic. (6) Base of posterior lateral
process less than one-half width of sternal plate at its middle.
CORACOID. (7) Entire proximal end straight above procoracoid
process. (8) Head narrowly rounded, moderately compressed, mod-
erately inflected. (9) Brachial tuberosity with slight dorsal and ventral
overhang. (10) Furcular facet obsolete. (11) Bicipital attachment
well developed, margin between it and head concave. (12) Shaft wide
proximally, distal portion of ventral surface usually convex medially.
(13) Procoracoid process with its apex rounded. (14) Dorsal inter-
muscular line swinging out to medial surface of shaft, distal half
sharply raised. (15) Ventral intermuscular line terminating at tip of
sterno-coracoid process. (16) Dorsal face deeply excavated distally,
without fossa. (17) Tubercle for ligamentum sterno-coracoideum dor-
sale usually well developed and rounded: (18) Internal distal angle
without distinct tubercle or ridge on dorsal face. (19) Sterno-cora-
coidal process with distal margin slightly oblique, not produced as a
sharply raised ridge, without dorsal groove; its tip directed laterally
and ending in a terminal knob. (20) Distal border of sternal facet
deeply concave.





BULLETIN FLORIDA STATE MUSEUM


SCAPULA. (21) Glenoid facet oval in dorsal view. (22) Dorsal de-
pression just media to glenoid facet obsolete to shallowly excavated.
(23) Bridge between acromion process and glenoid facet about one-
half width of glenoid facet. (24) Acromion process with apex rounded,
strongly deflected.
HUMERUS. (25) Pneumatic fossa a moderately long ellipse or oval-
oid, without inner shelf. (26) Median crest strongly developed, ele-
vated only at its middle. (27) Capital groove moderately excavated,
its internal margin moderately elevated. (28) Margin between head
and internal tuberosity moderately concave. (29) Head broader than
long in anconal view. (30) Ridge along medial border of fossa II
moderately swollen, elongate, its apex acute. (31) Fossa II well de-
veloped. (32) Latissimus ridge originating on lateral edge of shaft,
but swinging in onto anconal face proximally. (33) External tuberosity
moderately inflated. (34) Bicipital crest arising gently from shaft,
pointed, its distal border lacking a groove. (35) Deltoid crest mod-
erately developed, slightly inflected, its summit knoblike. (36) Entepi-
condyle produced slightly beyond level of internal condyle. (37) En-
tepicondylar prominence moderately produced, its pit on palmar face.
(38) Depression of M. brachialis anticus shallowly excavated. (39)
Distal border of external condyle rounded in anconal view.
ULNA. (40) Olecranon process strongly produced, its apex pointed.
(41) External cotyla moderately elongate, rounded throughout, its
distal border sloping gently into shaft, only slightly notched by proxi-
mal radial impression. (42) Impression for M. brachialis anticus exca-
vated, its borders distinct. (43) Height at middle of shaft 60 to 63
percent of height just distal to external cotyla. (44) Distal radial im-
pression obsolete. (45) Notch between carpal tuberosity and internal
condyle well developed. (46) External condyle moderately produced,
usually arising abruptly from shaft, usually rounded throughout.
CARPOMETACARPUS. (47) Metacarpal I usually moderately pointed
terminally. (48) Ridge separating anterior carpal fossa from excavated
area above pisiform process produced, extends well onto base of meta-
carpal I. (49) Pisiform process oval or ovaloid in internal view, slightly
reflected dorsad, thus usually below, occasionally at level of dorsal rim
of inner carpal trochlea. (50) Ligamental attachment of pisiform proc-
ess produced and moderately swollen proximally, flattened distally,
proximal portion connected to pisiform process by bony bridge. (51)
Internal carpal trochlea with proximal border shallowly concave, its
ventral border without notch. (52) Intermetacarpal tubercle not reach-
ing or barely extending to metacarpal III, distally placed with small
space between its posterior border and junction of metacarpals II


Vol. 6





HOLMAN: OSTEOLOGY OF QUAILS


and III. (53) Tubercle on internal proximal face of metacarpal III
elongate and moderately produced. (54) One obsolete to weakly de-
veloped tubercle on external proximal face of metacarpal III. (55)
Metacarpal III slightly bowed.
PELVIS. (56) Width of pelvis through antitrochanters 50 to 54 per-
cent length of innominate from anterior edge of ilium to ischial angle.
(57) Pectineal process obsolete, only slightly produced beyond acetabu-
lar border as a minute point. (58) Depression on medial side of pec-
tineal process obsolete, without foramen. (59) Posterior iliac crest
with prominence strongly produced from just posterior to ilio-ischiac
fenestra. (60) No ridge or prominence ventral to posterior iliac crest.
(61) Dorsal face of postacetabular ilium narrow anteriad, further nar-
rowing gently posteriad to form pointed apex of much elongated, nar-
row, dorsal roof of posterior process. (62) Caudal face of postacetabu-
lar ilium, obsolete, narrowly triangular, horizontal in position, form-
ing ventral floor of posterior process. (63) Lateral face of postacetab-
ular ilium with its posterodorsal apex much pointed, forming much
elongated lateral wall of posterior process. (64) Renal bar broad, bar,
but not posterior process, excavated by renal depression. (65) Ischium
at angle of about 145 degrees to ventral edge of posterior iliac crest.
(66) Width through transverse processes of fourth synsacral vertebra
67 to 71 percent of length of synsacrum from centrum of first synsacral
vertebra through transverse process of fourth synsacral vertebra.
FEMUR. (67) Distal width 17 to 19 percent of length. (68) Head
moderately enlarged, neither bent dorsally nor rotated posteriorly.
(69) Neck short. (70) Iliac facet without compression, posterior border
convex. (71) Dorsal crest of trochanter depressed and deflected, its
dorsal border flat. (72) Trochanteric ridge 23 to 25 percent length of
femur. (73) Distal depth 75 to 79 percent of distal width. (74) In-
ternal condyle moderately wide, its anterior border usually weakly
produced throughout. (75) External condyle compressed, rounded
anteriorly.
TIBIOTARSUS. (76) Depth of inner cnemial crest 37 to 46 percent of
its length. (77) Outer cnemial crest usually straight. (78) Interarticu-
lar tubercle weakly produced. (79) Area between external articular
surface and outer cnemial crest moderately concave in dorsal view.
(80) Posterior margin between external and internal articular surface
with distinct notch. (81) Area just anterior to ridge for proximal in-
ternal ligamental attachment without deeply excavated depression.
(82) Internal ligamental prominence on internal condyle moderately
produced. (83) Face of internal condyle moderately excavated, its






BULLETIN FLORIDA STATE MUSEUM


border moderately elevated. (84) Dorsal border of supratendinal
bridge straight.
TARSOMETATARSUS. (85) Distal width 21 to 22 percent of length.
(86) Proximal depth 95 to 98 percent of proximal width. (87) Inter-
cotylar prominence moderately deflected, its apex pointed. (88) Pos-
terior intercotylar area usually moderately excavated as a triangular
depression. (89) Area surrounding posterior opening of inner proximal
foramen moderately excavated. (90) Hypotarsus with 2 well-devel-
oped and 1 obsolete calcaneal ridges, 2 calcaneal grooves, and 2 closed
calcaneal canals. (91) Anterior face of shaft usually moderately exca-
vated by anterior metatarsal groove. (92) Depth of middle trochlea
46 to 58 percent of distal width. (93) Trochlea for digit II moderately
inflected. (94) Wing of trochlea for digit IV depressed dorsoventrally,
thus entire trochlea much deeper than long.
INTERMEMBRAL PROPORTIONS. (95) Wing/leg 78 to 81 percent.
(96) Ulna/humerus 93 to 94 percent. (97) Tarsometatarsus/femur 70
to 73 percent. (98) Tarsometatarsus/tibiotarsus 53 to 56 percent. (99)
Humerus/femur 95 to 101 percent. (100) Humerus/tibiotarsus 72 to
75 percent. (101) Ulna/femur 88 to 93 percent. (102) Ulna/tibiotarsus
68 to 71 percent. (103) Ulna/tarsometatarsus 126 to 133 percent.
(104) Carpometacarpus/femur 50 to 52 percent. (105) Carpometacar-
pus/tibiotarsus 38 to 40 percent. (106) Carpometacarpus/tarsometa-
tarsus 71 to 74 percent. (107) Tarsometatarsus/humerus 70 to 74 per-
cent. (108) Ilium/sternum 72 to 76 percent. (109) Height of sternal
carina/ilium 38 to 41 percent.

Rhynchortyx OGILVIE-GRANT, 1893
The monotypic genus Rhynchortyx occurs in the tropical lowlands
from Honduras to Ecuador. Measurements of R. cinctus (Salvin, 1876)
are given in table 1.
ROSTRUM. (1) Short and deep. (2) Nasal fossae round.
STERNUM. (3) Height of carina through sternal plate 35 percent
length of sternum. (4) Dorsal lip of coracoidal sulcus projected only
slightly beyond ventral lip. (5) Depressions on anterior portion of ster-
nal plate deep and pneumatic. (6) Base of posterior lateral process less
than one-half width of sternal plate at its middle.
CORACOID. (7) Entire proximal end straight above procoracoid
process. (8) Head broadly rounded, much depressed, moderately in-
flected. (9) Brachial tuberosity with slight dorsal and obsolete ven-
tral overhang. (10) Furcular facet obsolete. (11) Bicipital attachment
weakly developed, margin between it and head straight. (12) Shaft


Vol. 6






BULLETIN FLORIDA STATE MUSEUM


CARPOMETACARPUS. (47) Metacarpal I moderately pointed ter-
minally. (48) Ridge separating anterior carpal fossa from excavated
area above pisiform process produced, extending well onto base of met-
acarpal I. (49) Pisiform process elliptical in internal view, slightly
reflected dorsad, thus below level of dorsal rim of inner carpal trochlea.
(50) Ligamental attachment of pisiform process weakly produced and
only slightly swollen proximally, flattened distally, proximal portion
connected to pisiform process by bony bridge. (51) Internal carpal
trochlea with proximal border shallowly concave, its ventral border
without notch. (52) Intermetacarpal tubercle extending to but not
ankylosing with metacarpal III, distally placed with a small space be-
tween posterior border and junction of metacarpals II and III. (53)
Tubercle on internal proximal face of metacarpal III elongate and
moderately produced. (54) One obsolete tubercle on external proximal
face of metacarpal III. (55) Metacarpal III slightly bowed.
PELVIS. (56) Width of pelvis through antitrochanters 59 percent
length of innominate from anterior edge of ilium to ischial angle.
(57) Pectineal process obsolete, only slightly produced beyond ace-
tabular border as a minute point. (58) Depression on medial side of
pectineal process obsolete, without foramen. (59) Posterior iliac crest
with prominence weakly produced from just posterior to ilio-ischiac
fenestra. (60) Without ridge or prominence ventral to posterior iliac
crest. (61) Dorsal face of postacetabular ilium moderately broadened
anteriad, narrowing gently posteriad to form truncated apex of mod-
erately long, narrow, dorsal roof of posterior process. (62) Caudal
face of postacetabular ilium obsolete, rectangular, horizontal in posi-
tion, forming ventral floor of posterior process. (63) Lateral face of
postacetabular ilium with posterodorsal apex truncated, forming mod-
erately elongate lateral wall of posterior process. (64) Renal bar
broad, bar slightly excavated, but posterior process unexcavated by
renal depression. (65) Ischium at angle of about 160 degrees to ven-
tral edge of posterior iliac crest. (66) Width through transverse proc-
esses of fourth synsacral vertebra 92 percent length of synsacrum from
centrum of first synsacral vertebra through transverse process of
fourth synsacral vertebra.
FEMUR. (67) Distal width 18 percent of length. (68) Head mod-
erately enlarged, bent dorsally, but without posterior rotation. (69)
Neck moderately long. (70) Iliac facet without compression, posterior
border convex. (71) Dorsal crest of trochanter obsolete, its dorsal
border flat. (72) Trochanteric ridge 12 percent length of femur. (73)
Distal depth 71 percent of distal width. (74) Internal condyle mod-
erately wide, its anterior border much produced throughout. (75)
External condyle compressed, rounded anteriorly.


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HOLMAN: OSTEOLOGY OF QUAILS


TIBIOTARSUS. (76) Depth of inner cnemial crest 75 percent of its
length. (77) Outer cnemial crest decurved. (78) Interarticular tu-
bercle moderately produced. (79) Area between external articular
surface and outer cnemial crest moderately concave in dorsal view.
(80) Posterior margin between external and internal articular surface
with distinct notch. (81) Area just anterior to ridge for proximal in-
ternal ligamental attachment without deeply excavated depression.
(82) Area of internal ligamental attachment on internal condyle not
produced as a prominence. (83) Face of internal condyle weakly ex-
cavated, its border weakly elevated. (84) Dorsal border of supra-
tendinal bridge oblique.
TARSOMETATARSUS. (85) Distal width 16 percent of length. (86)
Proximal depth 95 percent of proximal width. (87) Intercotylar prom-
inence depressed, without deflection, its apex broadly rounded. (88)
Posterior intercotylar area unexcavated. (89) Area surrounding pos-
terior opening of inner proximal foramen shallowly excavated. (90)
Hypotarsus with 3 well-developed calcaneal ridges, 2 calcaneal
grooves, and 1 closed calcaneal canal. (91) Anterior face of shaft ex-
tensively excavated by anterior metatarsal groove. (92) Depth of
middle trochlea 56 percent of distal width. (93) Trochlea for digit II
only slightly inflected. (94) Wing of trochlea for digit IV depressed
dorsoventrally, thus entire trochlea much deeper than long.
INTERMEMBRAL PROPORTIONS. (95) Wing/leg 67 percent. (96)
Ulna/humerus 98 percent. (97) Tarsometatarsus/femur 88 percent.
(98) Tarsometatarsus/tibiotarsus 63 percent. (99) Humerus/femur 85
percent. (100) Humerus/tibiotarsus 61 percent. (101) Ulna/femur 87
percent. (102) Ulna/tibiotarsus 62 percent. (103) Ulna/tarsometatar-
sus 99 percent. (104) Carpometacarpus/femur 48 percent. (105)
Carpometacarpus/tibiotarsus 34 percent. (106) Carpometacarpus/
tarsometatarsus 55 percent. (107) Tarsometatarsus/humerus 103 per-
cent. (108) Ilium/sternum 68 percent. (109) Height of sternal carina/
ilium 52 percent.

SUMMARY OF GENERIC CHARACTERS
Important osteological characters of the New World quails are of
four main types: (1) Characters of the pelvis that tend to divide the
New World quails into two major groups. An example is the poorly
developed, slender, renal bar in Dendrortyx, Philortyx, Oreortyx, Calli-
pepla, Colinus, and Lophortyx, and the well-developed, broad renal
bar in Odontophorus, Dactylortyx, Cyrtonyx, and Rhynchortyx. (2)
Primitive characters that are apparently holdovers from less highly





BULLETIN FLORIDA STATE MUSEUM


evolved groups (Holman, MS). An example is the indistinct fossa II
of the humerus in Dendrortyx and Odontophorus. (3) Characters that
are unique in certain genera, as the short, deep rostrum with round
nasal fossae in Rhynchortyx. (4) Characters that are not unique, but
may be used in combination with other characters for generic defini-
tion.
When a character occurs in most but not all individuals of the
genus, the condition is modified by "usually." Characters that hold
in some genera, but vary specifically or show much individual varia-
tion in other genera, are listed as "variable." Characters that show
much individual variation in all genera, and ratios that would prob-
ably overlap in larger series, are omitted from the following section.

ROSTRUM. The rostrum is relatively long and shallow and the nasal
fossae are elliptical in most New World quails. In Rhynchortyx the
rostrum is much shorter and deeper, and its nasal fossae are round.

STERNUM. The dorsal lip of the coracoidal sulcus projects well be-
yond the ventral lip in most genera. In Dactylortyx the dorsal lip
projects only slightly beyond the ventral lip, and in Rhynchortyx the
dorsal lip lies even with or projected only slightly beyond the ventral
lip. The character is variable in Cyrtonyx and Odontophorus.
The depressions on the anterior portion of the sternal plate are
deep and very pneumatic in Rhynchortyx, deep and moderately pneu-
matic in Oreortyx, deep and slightly pneumatic in Philortyx, shal-
low and slightly pneumatic in Dendrortyx, usually shallow and non-
pneumatic in Colinus, shallow and nonpneumatic in Cyrtonyx, shal-
low or obsolete and nonpneumatic in Dactylortyx, obsolete and non-
pneumatic in Odontophorus, and variable in Callipepla and Lophortyx.
The base of the posterior lateral process is narrow and less than
one-half the width of the sternal plate at its middle in most genera,
but it is wide and more than one-half the width of the sternal plate
at its middle in Dendrortyx.

CORACOID. The entire proximal end is straight above the level of
the procoracoid process in most species, but it is bent ventrad above
the level of the procoracoid process in Dendrortyx.
The head is narrowly rounded and moderately compressed in most
genera, but it is pointed and much compressed in Philortyx, broadly
rounded and much depressed in Dendrortyx and Rhynchortyx, and
broadly rounded and moderately depressed in Oreortyx.
The head is moderately inflected in Callipepla, Lophortyx, Dac-
tylortyx, Cyrtonyx, and Rhynchortyx, and much inflected in Dendror-


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HOLMAN: OSTEOLOGY OF QUAILS


tyx, but it is moderately reflected in Oreortyx, much reflected in Philor-
tyx, and variable in Colinus and Odontophorus.
The brachial tuberosity has a slight dorsal and ventral overhang
in most genera, but it has a moderate dorsal and slight ventral over-
hang in Dendrortyx, a slight dorsal and obsolete ventral overhang in
Rhynchortyx, an obsolete dorsal and slight ventral overhang in Philor-
tyx, and is variable in Odontophorus.
The furcular facet is obsolete in Rhynchortyx and Cyrtonyx, weakly
inflated in Dendrortyx, Philortyx, and Lophortyx, moderately inflated
in Oreortyx, Callipepla, and Dactylortyx, much inflated in Odontoph-
orus, and variable in Colinus.
The bicipital attachment is well developed, with the margin be-
tween it and the head concave in most genera, but in Rhynchortyx
it is weakly developed and with the margin between it and the head
straight.
The procoracoid process has its apex rounded in most genera,
but it is acute in Philortyx and Callipepla, and it is variable in Colinus.
The dorsal intermuscular line swings out to the medial surface of
the shaft in most genera, but the line swings out to the lateral border
of the shaft in Dendrortyx. The distal half of the dorsal intermuscu-
lar line is sharply raised in most forms, but the sharply raised portion
is confined to the distal three-fourths in Dactylortyx, the distal two-
thirds or three-fourths in Odontophorus, and only the distal one-fifth
in Dendrortyx.
The ventral intermuscular line terminates at the tip of the sterno-
coracoidal process in most genera. It terminates in the middle of
the distal border of the sterno-coracoidal process in Dendrortyx.
The dorsal face is weakly excavated distally in Dendrortyx, Philor-
tyx, Oreortyx, Callipepla, and Rhynchortyx, moderately excavated dis-
tally in Colinus, deeply excavated distally in Cyrtonyx and Dactylor-
tyx, and variable in Lophortyx. and Odontophorus. The dorsal face
lacks a distinct fossa in most genera, but has a distinct, round, deep
fossa below the tubercle for the ligamentum sterno-coracoideum dor-
sale and above the external end of the sternal facet in Dendrortyx.
The tubercle for the ligamentum sterno-coracoideum dorsale is
obsolete and irregular in shape in Rhynchortyx and Odontophorus,
obsolete and rounded in Dendrortyx and Callipepla, well developed
but irregular in shape in Oreortyx, well developed and rounded in
Philortyx, and variable in the other genera.
The internal distal angle is without a distinct tubercle or ridge in
most Odontophoridae. It has a distinct but irregular tubercle that






BULLETIN FLORIDA STATE MUSEUM


extends obliquely to above the lateral end of the sternal facet in Oreor-
tyx and Callipepla. In Philortyx and Rhynchortyx it has a distinct
oblique ridge; this structure is weakly developed in the latter genus.
The sterno-coracoidal process has its distal margin slightly oblique,
neither produced as a sharply raised ridge, nor grooved dorsally, and
its tip is directed laterally and possesses a terminal knob in most New
World quails. Philortyx differs in that the distal margin of the sterno-
coracoidal process is moderately oblique, Rhynchortyx in that the
distal margin is moderately oblique and produced as a sharply raised
ridge, and Dendrortyx in that the distal margin is strongly oblique,
grooved dorsally, with the tip of the sterno-coracoidal process di-
rected proximally and without a terminal knob.
The distal border of the sternal facet is deeply concave in Oreor-
tyx, Rhynchortyx, and Dactylortyx, usually deeply concave in Colinus,
shallowly concave in Philortyx, very shallowly concave in Dendrortyx,
and variable in the other genera.

SCAPULA. The glenoid facet is oval in dorsal view in most forms,
but it is elliptical in dorsal view in Dendrortyx and variable in Odon-
tophorus.
The bridge between the acromion process and glenoid facet is
about one-half the width of the glenoid facet in most genera, but
about one-third the width in Philortyx, and variable in Colinus, Loph-
ortyx, and Odontophorus.
The acromion process has its apex rounded in most genera, but it
is acute in Dendrortyx and Oreortyx, and truncated in Rhynchortyx.
The amount of deflection of the acromion process shows much intra-
generic variation, but it is much more strongly deflected in Cyrtonyx
than in the other forms.
The shape of the pneumatic fossa shows much variation within
genera, but only one species, Odontophorus gujanensis, has a perfo-
rated inner shelf that extends from the external bicipital surface to
the medial bar.
The median crest is strongly to very strongly developed in Odon-
tophorus, strongly developed in Dendrortyx, Oreortyx, Cyrtonyx,
Rhychortyx, and Dactylortyx, moderately to strongly developed in
Colinus, usually weakly developed in Callipepla, weakly developed
in Philortyx, and variable in Lophortyx. The median crest is elevated
only at its middle in most genera, but is elevated throughout in
Dendrortyx, Philortyx, and Odontophorus, and variable in Lophortyx.
The capital groove is moderately excavated and its internal mar-
gin is moderately elevated in most New World quails, but it is deeply


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HOLMAN: OSTEOLOGY OF QUAILS


excavated with its internal margin strongly elevated in Dendrortyx
and Philortyx.
The margin between the head and internal tuberosity is mod-
erately concave in most genera, but is only slightly concave in Den-
drortyx.
The head is broader than long in most genera, but it is about as
long as broad in Dendrortyx and Philortyx and is variable in Lophor-
tyx.
The ridge along the medial border of fossa II is moderately swol-
len in Callipepla, Lophortyx, Odontophorus, Cyrtonyx, and Rhynchor-
tyx, slightly to moderately swollen in Colinus, slightly swollen in
Dendrortyx and Philortyx, and much swollen in Oreortyx and Dac-
tylortyx. The ridge along the medial border of fossa II is elongate and
pointed in most genera, but it is short and rounded in Dactylortyx.
Fossa II is well developed in most forms but is obsolete in Den-
drortyx and Odontophorus.
The latissimus ridge originates on the lateral edge of the shaft
and swings onto the anconal face proximally in most New World
quails, but the ridge extends along the lateral edge of the shaft
throughout its length in Dendrortyx.
The external tuberosity is moderately inflated in most forms, but
it is much inflated in Oreortyx and variable in Odontophorus.
The bicipital crest arises gently from the shaft in most odonto-
phorids but arises abruptly from the shaft in Rhynchortyx. The bi-
cipital crest is pointed in Philortyx, Oreortyx, Cyrtonyx, and Dactylor-
tyx, usually narrowly rounded in Colinus, narrowly rounded in Calli-
pepla and Lophortyx, and rounded in Dendrortyx, Rhynchortyx, and
Odontophorus. The bicipital crest lacks a groove in its distal margin
in most forms, but has a shallow groove in Rhynchortyx.
The deltoid crest is moderately developed in most forms, but it
is much developed in Dendrortyx, and variable in Odontophorus.
The deltoid crest is slightly inflected in most forms but is much in-
flected in Dendrortyx. The summit of the deltoid crest is knoblike in
most, but it is usually flattened in Lophortyx, and variable in Colinus.
The entepicondyle is at the level of the internal condyle in Dend-
rortyx, Oreortyx, Callipepla, and Rhynchortyx, usually at the same
level in Colinus and Dactylortyx, and produced slightly beyond the
internal condyle in Philortyx, Lophortyx, Odontophorus, and Cyr-
tonyx.
The entepicondylar prominence is weakly produced in Dendrortyx,
moderately produced in Philortyx, Oreortyx, Callipepla, Colinus, Loph-
ortyx, and Cyrtonyx, and much produced in Odontophorus, Dactylor-






BULLETIN FLORIDA STATE MUSEUM


tyx, and Rhynchortyx. The pit of the entepicondylar prominence is
placed on the internal face of the humerus in most forms, but it is on
the palmar face in Callipepla, Cyrtonyx, and Rhynchortyx.
The depression for the M. brachialis anticus is deeply excavated in
Dendrortyx and Philortyx, usually moderately excavated in Colinus,
moderately excavated in Lophortyx and Odontophorus, usually shal-
lowly excavated in Callipepla, shallowly excavated in Oreortyx, Cyr-
tonyx, and Dactylortyx, and represented by an unexcavated scar in
Rhynchortyx.
The distal border of the external condyle is usually rounded in
anconal view, but it is flat in Dendrortyx.

ULNA. The olecranon process is usually strongly produced, but is
weakly produced in Rhynchortyx. The apex of the olecranon process
is pointed in most forms, but it is rounded in Dendrortyx and Rhyn-
chortyx, and truncated in Dactylortyx.
The external cotyla is moderately elongate in most genera, but it
is much elongated in Oreortyx, and shortened in Dendrortyx. The
external cotyla is rounded throughout, and its distal border slopes
gently into the shaft in most forms, but it is flattened dorsally with its
distal border sloping precipitously into the shaft in Dendrortyx, and
it is variable in Odontophorus. The external cotyla is slightly notched
by the proximal radial impression in Oreortyx, Callipepla, Colinus,
Cyrtonyx, and Dactylortyx, usually slightly notched in Colinus, deeply
notched in Dendrortyx, Philortyx, and Rhynchortyx, and is variable
in Odontophorus.
The impression for the M. brachialis anticus is excavated with its
borders distinct in most genera, but it is represented by an unexca-
vated scar with indistinct borders in Philortyx.
The height at the middle of the shaft usually ranges from 60 to 72
percent of the height of the shaft just distal to the external cotyla,
with much overlap of ranges between genera. This ratio is 83 per-
cent in Dendrortyx and 85 percent in Philortyx.
The distal radial impression is obsolete in most genera, well ex-
cavated in Dendrortyx, and variable in Odontophorus.
The notch between the carpal tuberosity and internal condyle is
well developed in most forms, but it is obsolete in Dendrortyx.
The external condyle is well produced in Philortyx, Callipepla,
Colinus, Lophortyx, and Odontophorus, moderately produced in Oreor-
tyx, Cyrtonyx, and Dactylortyx, and weakly produced in Dendrortyx
and Rhynchortyx. The external condyle arises abruptly from the
shaft in most forms, usually arises abruptly from the shaft in Cyrtonyx,


Vol. 6






HOLMAN: OSTEOLOGY OF QUAILS


but arises gently from the shaft in Oreortyx, Lophortyx, and Rhynchor-
tyx. The external condyle is rounded throughout in most genera,
usually rounded in Cyrtonyx and Dactylortyx, but has its ventral bor-
der flattened in Dendrortyx and Philortyx.

CARPOMETACARPUS. Metacarpal I is moderately pointed terminally
in Dendrortyx, Philortyx, Callipepla, Odontophorus, and Rhynchortyx,
usually moderately pointed terminally in Cyrtonyx, sharply pointed
terminally in Oreortyx, usually rounded terminally in Dactylortyx,
and variable in Colinus and Lophortyx.
The ridge that separates the anterior carpal fossa from the exca-
vated area above the pisiform process is produced and extends well
onto the base of metacarpal I in most odontophorids, but it is weak
and extends only slightly onto the base of metacarpal I in Dendrortyx.
The pisiform process is ovaloid in internal view in Oreortyx, Lo-
phortyx, Dactylortyx, and Odontophorus, elliptical in Rhynchortyx,
round in Dendrortyx and Philortyx, and variable in Callipepla, Colinus,
and Cyrtonyx. The pisiform process is slightly reflected dorsad and
thus is below the level of the dorsal rim of the inner carpal trochlea
in Oreortyx, Rhynchortyx, Dactylortyx, and Odontophorus, slightly
reflected dorsad and usually below, but occasionally at the level of the
inner carpal trochlea in Callipepla, Colinus, and Cyrtonyx, moderately
reflected dorsad and thus produced slightly above the dorsal rim of
the inner carpal trochlea in Philortyx, much reflected dorsad and pro-
duced well above the dorsal rim of the inner carpal trochlea in Dend-
rortyx, and variable in Lophortyx.
The ligamental attachment of the pisiform process is produced
and much swollen proximally and flattened distally in most species.
Odontophorus differs in that the ligamental attachment is moderately
or much swollen proximally, Cyrtonyx in that it is moderately swollen
proximally, and Rhynchortyx in that it is weakly produced and only
slightly swollen proximally, and Dendrortyx in that the ligamental
attachment of the pisiform process is produced throughout and mod-
erately swollen proximally and distally. The proximal portion of the
ligamental attachment of the pisiform process is connected to the pisi-
form process by a bony bridge in most genera; the attachment is
usually connected by a bony bridge in Colinus, separated from the
pisiform process by a shallow sulcus in Dendrortyx, and variable in
Odontophorus.
The proximal border of the internal carpal trochlea is shallowly
concave in Dendrortyx, Philortyx, Rhynchortyx, Cyrtonyx, and Dac-
tylortyx, distinctly concave in Oreortyx, usually slightly notched in






BULLETIN FLORIDA STATE MUSEUM


Lophortyx and Odontophorus, deeply notched in Callipepla, and vari-
able in Colinus. The ventral border of the internal carpal trochlea
is without a notch in most genera, but it has a slight notch in Dendror-
tyx and Oreortyx.
The intermetacarpal tubercle extends to or slightly beyond meta-
carpal III without ankylosing with it in most New World quails; it
extends to and sometimes ankyloses with metacarpal III in Lophortyx,
extends to and ankyloses with metacarpal III in Dendrortyx, and does
not reach or barely extends to metacarpal III in Cyrtonyx. The inter-
metacarpal tubercle is relatively distally placed and has a small space
between its posterior border and junction of metacarpals II and III
in most genera, but it is more proximally placed and with only a
minute space between its posterior border and junction of metacarpals
II and III in Dactylortyx.
The tubercle on the internal proximal surface of metacarpal
III is obsolete in most forms, but it is elongate and weakly produced
in Philortyx, and elongate and moderately produced in Oreortyx, Cyr-
tonyx, and Rhynchortyx.
The external proximal face of metacarpal III has a single strongly
developed tubercle in Oreortyx, Callipepla, Colinus, Lophortyx, and
Odontophorus, a single moderately developed tubercle in Philortyx
and Dactylortyx, a single obsolete to weakly developed tubercle in
Cyrtonyx, a single obsolete tubercle in Rhynchortyx, and one mod-
erately developed and one obsolete tubercle in Dendrortyx.
Metacarpal III is slightly bowed in most forms, but is moderately
bowed in Dendrortyx.

PELVIS. The pelvis of Dendrortyx, Philortyx, and Colinus is relatively
narrow, and the ratio of the width of the pelvis through the anti-
trochanters divided by the length of the innominate from the anterior
edge of the ilium to the ischial angle ranges from 44 to 51 percent.
The pelvis of Oreortyx, Callipepla, and Rhynchortyx is relatively wide,
the ratio being 59 percent in Rhynchortyx, 58 to 62 percent in Calli-
pepla, and 60 to 63 percent in Oreortyx. The remaining genera are
poorly separable on this character.
The pectineal process is moderately produced beyond the acetabu-
lar border with its apex pointed in Philortyt, Oreortyx, Callipepla, and
Lophortyx, usually pointed in Colinus, but rounded in Dendrortyx.
The pectineal process is only slightly produced beyond the acetabular
border as a minute point in Odontophorus, Cyrtonyx, and Rhynchortyx,
and in Dactylortyx it is usually absent or at most, represented by a
minute point.


Vol. 6






HOLMAN: OSTEOLOGY OF QUAILS


The depression on the medial side of the pectineal process is deeply
excavated and perforated by a foramen in Dendrortyx, deeply exca-
vated and without a foramen in Philortyx, and shallowly excavated and
without a foramen in Oreortyx, Callipepla, Colinus, and Lophortyx.
The depression on the medial side of the pectineal process is obsolete
and without a foramen in Odontophorus, Cyrtonyx, and Rhynchortyx,
and absent in Dactylortyx.
The posterior iliac crest lacks a prominence in most genera. A
prominence is weakly produced just posterior to the ilio-ischiac fen-
estra in Rhynchortyx, moderately produced in Dactylortyx, and strong-
ly produced in Cyrtonyx.
A ridge originates on the posterior iliac crest at the level of the
ilio-ischiac fenestra, continues ventrad and posteriad towards the pos-
terior border of the ilium, and has a prominence arising from its
posterior portion in most species. The prominence arises sharply in
Dendrortyx, Philortyx, Lophortyx, and Colinus, but it is weak in
Oreortyx and Callipepla. The ridge and prominence are absent in
Dactylortyx, Cyrtonyx, and Rhynchortyx. Odontophorus is variable
in this character.
The dorsal face of the postacetabular ilium is moderately broad
anteriad and narrows abruptly posteriad to form the acute apex of the
short, broad, dorsal roof of the posterior process in Dendrortyx, Philor-
tyx, Callipepla, Colinus, and Lophortyx. It is much broadened an-
teriad, and narrows abruptly posteriad to form the acute apex of the
short, broad, dorsal roof of the posterior process in Oreortyx. In
Dactylortyx and Rhynchortyx the dorsal face of the postacetabular
ilium is moderately broadened anteriad and narrows abruptly pos-
teriad to form the truncated apex of the moderately long, narrow, dor-
sal roof of the posterior process. Cyrtonyx has the dorsal face of the
postacetabular ilium narrow anteriad, and it gradually further nar-
rows posteriad to form the pointed apex of the much elongated, nar-
row, dorsal roof of the posterior process. Odontophorus is variable
in this character.
The caudal face of the postacetabular ilium is enlarged, triangular,
and vertical in position, and its posterodorsal apex forms the postero-
medial wall of the posterior process in Dendrortyx, Philortyx, Calli-
pepla, Colinus, and Lophortyx. Oreortyx differs only in that the caudal
face is broadly triangular. The caudal face of the postacetabular
ilium is obsolete, horizontal in position, and forms the ventral floor of
the posterior process in Odontophorus, Dactylortyx, Cyrtonyx, and
Rhynchortyx. It is rectangular in shape in Dactylortyx and Rhynchor-
tyx, triangular in Odontophorus, and narrowly triangular in Cyrtonyx.






BULLETIN FLORIDA STATE MUSEUM


The lateral face of the postacetabular ilium generally has an acute
posterodorsal apex, but it is much pointed in Cyrtonyx and truncated
in Rhynchortyx. It forms the shortened lateral wall of the posterior
process in most genera, but is moderately elongate in Rhynchortyx,
elongate in Dactylortyx, moderately to much elongated in Odonto-
phorus, and much elongated in Cyrtonyx.
The renal bar is slender, and both the bar and the posterior process
are excavated by the renal depression in Dendrortyx, Callipepla,
Colinus, and Lophortyx. It is moderately slender in Philortyx, obso-
lete in Oreortyx, and both the renal bar and the posterior process are
excavated by the renal depression in these genera. The renal bar is
broad; the bar, but not the posterior process, is excavated by the renal
depression in Odontophorus, Dactylortyx, and Cyrtonyx. The bar is
broad but only slightly excavated in Rhynchortyx.
The ischium lies at an angle of about 160 degrees to the ventral
edge of the posterior iliac crest in most genera, but it lies at an angle
of about 145 degrees to the ventral edge of the posterior iliac crest in
Cyrtonyx.
The synsacrum of Dendrortyx is long and narrow with a 55 percent
ratio for the width through the transverse processes of the fourth
synsacral vertebra divided by the length of the synsacrum from the
centrum of the first synsacral vertebra through the transverse process
of the fourth synsacral vertebra. The synsacra of Rhynchortyx and
Dactylortyx are much shorter and wider, with the ratio 88 to 93 per-
cent in Dactylortyx and 92 percent in Rhynchortyx. The other genera
have the synsacrum shorter and wider than Dendrortyx, and longer
and narrower than Rhynchortyx and Dactylortyx, but are poorly sep-
arable among themselves.

FEMUR. The head is moderately enlarged in most forms, but it is
much enlarged in Dactylortyx and is variable in Lophortyx. The head
is neither bent dorsally nor rotated posteriorly in most genera, but it
is bent dorsally but without posterior rotation in Rhynchortyx, and
bent dorsally and rotated posteriorly in Dendrortyx.
The neck is only moderately long in most forms, but it is long in
Dendrortyx, and short in Cyrtonyx.
The iliac facet is without compression, and its dorsal border is flat
in most genera, but it is compressed with its posterior border straight
in Dendrortyx.
The dorsal crest of the trochanter is compressed and reflected with
its dorsal border rounded in Philortyx, Oreortyx, Callipepla, Dactylor-
tyx, usually compressed and reflected with the dorsal border rounded


Vol. 6





HOLMAN: OSTEOLOGY OF QUAILS


in Lophortyx, depressed and deflected with its dorsal border flat in
Dendrortyx, Odontophorus, and Cyrtonyx, obsolete and flattened in
Rhynchortyx, and variable in Colinus.
The trochanteric ridge is normally about 20 to 25 percent of the
length of the femur, but the ratio is only 12 percent in Rhynchortyx.
The distal depth is about 80 to 90 percent of the distal width in
most genera, but the ratio is 71 percent in Rhynchortyx, and 97 per-
cent in Philortyx.
The internal condyle is moderately wide and has its anterior bor-
der much produced throughout in most forms. Cyrtonyx differs only
in that the anterior border of the internal condyle is weakly produced
throughout. Dactylortyx differs in that the internal condyle is nar-
row and usually has its anterior border weakly produced throughout,
whereas in Philortyx the anterior border of the condyle is very wide,
and its border is moderately produced throughout.
The external condyle is ordinarily compressed and rounded an-
teriorly, but it is depressed and flattened in Dendrortyx.

TIBIOTABsUS. The depth of the inner cnemial crest is usually well
over 50 percent of its length, but the ratio is 42 to 48 percent in Dac-
tylortyx, 45 percent in Philortyx, and 37 to 46 percent in Cyrtonyx.
The outer cnemial crest is straight in Oreortyx, Odontophorus, and
Dactylortyx, usually straight in Callipepla, Colinus, and Cyrtonyx,
decurved in Dendrortyx, Philortyx, and Rhynchortyx, and variable in
Lophortyx.
The interarticular tubercle is weakly produced in Philortyx, Dacty-
lortyx, Cyrtonyx, moderately produced in Dendrortyx, Callipepla,
Lophortyx, and Rhynchortyx, usually moderately produced in Colinus,
moderately or strongly produced in Odontophorus, and strongly pro-
duced in Oreortyx.
The area between the external articular surface and outer cnemial
crest, normally moderately concave in dorsal view, is moderately or
deeply concave in Odontophorus and very deeply concave in Dactylor-
tyx.
The posterior margin between the external and internal articular
surface has a distinct notch in most genera, but it is weakly notched
in Dendrortyx.
In most species the area just anterior to the ridge for the proximal
internal ligamental attachment lacks the deeply excavated depression
present in Dendrortyx.
The area of the internal ligamental attachment on the internal
condyle is moderately produced as a prominence in most genera,





BULLETIN FLORIDA STATE MUSEUM


strongly produced in Dendrortyx, and lacks a prominence in Dactylor-
tyx and Rhynchortyx.
The face of the internal condyle is moderately excavated with its
border moderately elevated in most genera, but it is well excavated
with its border highly elevated in Dendrortyx, and weakly excavated
with a slightly elevated border in Rhynchortyx.
The dorsal border of the supratendinal bridge is straight in Philor-
tyx, CaUipepla, Dactylortyx, and Cyrtonyx, usually straight in Oreor-
tyx, Lophortyx, and Odontophorus, oblique in Dendrortyx and Rhyn-
chortyx, and variable in Colinus.
TARSOMETATARSUS. The intercotylar prominence is moderately de-
flected in most species, but it is much deflected in Philortyx, only
slightly deflected in Dactylortyx, and depressed and without deflection
in Rhynchortyx. The apex of the intercotylar prominence is pointed
in Dendrortyx, Philortyx, Lophortyx, Odontophorus, and Cyrtonyx,
usually pointed in Oreortyx, Callipepla, Colinus, and Dactylortyx, and
broadly rounded in Rhynchortyx.
The posterior intercotylar area is moderately excavated as a tri-
angular depression in Dendrortyx, Odontophorus, and Dactylortyx,
usually moderately excavated as a triangular depression in Cyrtonyx,
weakly excavated as a triangular depression in Philortyx, Colinus, and
Callipepla, usually weakly excavated as a triangular depression in
Lophortyx, unexcavated in Rhynchortyx, and usually unexcavated in
Oreortyx.
The area surrounding the posterior opening of the inner proximal
foramen is shallowly excavated in Oreortyx, Callipepla, Colinus, Lo-
phortyx, and Rhynchortyx, moderately excavated in Dactylortyx and
Cyrtonyx, deeply in Dendrortyx and Philortyx, and very deeply in
Odontophorus.
The hypotarsus has 2 well developed and 1 obsolete calcaneal
ridges in most genera, but there are 2 well developed and 1 weakly
developed calcaneal ridges in Dactylortyx, and 3 well developed cal-
caneal ridges in Rhynchortyx. The hypotarsus has 2 closed calcaneal
canals in most forms, but there is only 1 closed calcaneal canal in
Dendrortyx and Rhynchortyx, usually 1 in Oreortyx, and the condition
is variable in Odontophorus.
The anterior face of the shaft is extensively excavated by the an-
terior metatarsal groove in Dendrortyx, Philortyx, and Rhynchortyx,
moderately excavated by the anterior metatarsal groove in Callipepla,
usually moderately excavated in Colinus and Cyrtonyx, usually re-
duced in Lophortyx, reduced in Odontophorus, reduced or obsolete in
Dactylortyx, and reduced or absent in Oreortyx.


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HOLMAN: OSTEOLOGY OF QUAILS


The trochlea for digit II is only slightly inflected in most genera,
but is moderately inflected in Oreortyx, Dactylortyx, and Cyrtonyx.
The wing of the trochlea for digit IV is depressed dorsoventrally
with the entire trochlea much deeper than long in most genera, but
in Dendrortyx and Odontophorus the wing is depressed anteropos-
teriorly, and the entire trochlea is longer than deep.

INTERMEMBRAL PROPORTIONS. The ratio of the combined length of
the bones of the wing (humerus, ulna, carpometacarpus) divided by
the combined length of the bones of the leg (femur, tibiotarsus, tarso-
metatarsus) divides the New World quails into three groups. Dendror-
tyx has a relatively short wing with a ratio of 52 percent. Cyrtonyx
and Dactylortyx have relatively long wings; this ratio is 78 to 81 per-
cent in Cyrtonyx and 72 to 73 percent in Dactylortyx. The other forms
are intermediate; the ratio ranges from 60 to 69 percent, with consid-
erable overlap among genera.
The ulna is more than 95 percent of the length of the humerus in
Odontophorus, Dactylortyx, and Rhynchortyx. The other forms range
from 86 to 94 percent in this ratio with much overlap.
The tarsometatarsus is more than 85 percent of the length of the
femur in Dendrortyx and Rhynchortyx and 84 to 87 percent in Odon-
tophorus. The other forms range from 70 to 83 percent in this ratio
with much overlap among genera.
The tarsometatarsus is 63 percent or more of the length of the tibio-
tarsus in Dendrortyx, Odontophorus, and Rhynchortyx. The other
forms range from 53 to 61 percent in this ratio with considerable over-
lap among genera.
The humerus is 71 percent of the length of the femur in Dendror-
tyx, but 90 to 91 percent in Dactylortyx, and 95 to 100 percent in
Cyrtonyx. The other forms are intermediate with considerable over-
lap.
The humerus is 51 percent of the length of the tibiotarsus in
Dendrortyx and 72 to 75 percent in Cyrtonyx. The other forms are
intermediate with much overlap in the ratio among genera.
The ulna is 63 percent of the length of the femur in Dendrortyx,
87 percent in Rhynchortyx, 88 to 91 percent in Dactylortyx, and
89 to 93 percent in Cyrtonyx. The other forms are intermediate with
considerable overlap.
The ulna is 45 percent of the length of the tibiotarsus in Dendror-
tyx, 65 to 66 percent in Dactylortyx, and 68 to 71 percent in Cyrtonyx.
The others are intermediate with much overlap among genera.
The ulna is 72 percent of the length of the tarsometatarsus in






BULLETIN FLORIDA STATE MUSEUM


Dendrortyx and 126 to 133 percent in Cyrtonyx. The others are in-
termediate with considerable overlap among genera.
The carpometacarpus is 27 percent of the length of the tibiotarsus
in Dendrortyx, and 38 to 40 percent in Cyrtonyx. The other forms are
intermediate with much overlap.
The carpometacarpus is 42 percent of the length of the tarsometa-
tarsus in Dendrortyx, and 71 to 74 percent in Cyrtonyx. The other
forms are intermediate with much overlap among genera.
The tarsometatarsus is 124 percent of the length of the humerus in
Dendrortyx, but only 70 to 74 percent in Cyrtonyx. The others are
intermediate with considerable overlap.
The ilium is 89 percent of the length of the sternum in Dendror-
tyx, whereas this ratio ranges from 65 to 78 percent in the other forms,
with much overlap among genera.
The height of the sternal carina is 34 percent of the length of the
ilium in Dendrortyx, and 52 percent in Rhynchortyx. The other forms
are intermediate with much overlap.

MAJOR RELATIONSHIPS
The New World quails may be divided into two distinct groups on
the basis of striking differences in pelvic structures (figs. 1 and 2),
referred to here as the Dendrortyx group, and the Odontophorus
group. The genera in each of the two groups are:
Dendrortyx group Odontophorus group
Dendrortyx Odontophorus
Philortyx Dactylortyx
Oreortyx Cyrtonyx
Callipepla Rhynchortyx
Colinus
Lophortyx
Differences in the two basic pelvic types are: 1) In the Dendrortyx
type the pectineal process is moderately produced beyond the ace-
tabular border. In the Odontophorus type the pectineal process is
represented only by a minute point or is absent. 2) In the Dendrortyx
type the depression on the medial side of the pectineal process is dis-
tinct. The depression is obsolete or absent in the Odontophorus type.
3) In the Dendrortyx type the posterior iliac crest is without a prom-
inence. A prominence is usually produced from the posterior iliac
crest in the Odontophorus type. 4) In the Dendrortyx type a ridge
originates on the posterior iliac crest at the level of the ilio-ischiac
fenestra, continues ventrad and posteriad toward the posterior border
of the ilium, and has a prominence that arises from its posterior por-


Vol. 6





HOLMAN: OSTEOLOGY OF QUAILS


tion. In the Odontophorus type the ridge and prominence are absent,
except in one specimen of Odontophorus gujanensis. 5) The dorsal
face of the postacetabular ilium is relatively short and broad in the
Dendrortyx type. In the Odontophorus type the dorsal face is rela-
tively long and narrow. 6) In the Dendrortyx type the caudal face
of the postacetabular ilium is enlarged, triangular, vertical in position,
and its posterodorsal apex forms the posteromedial wall of the pos-
terior process. In the Odontophorus type the caudal face of the post-
acetabular ilium is obsolete, variable in shape, horizontal in position,
and forms the ventral floor of the posterior process. 7) The lateral
face of the postacetabular ilium is relatively short in the Dendrortyx
type. In the Odontophorus type the lateral face is relatively long.
8) In the Dendrortyx type the renal bar is poorly developed or obsolete,
always slender, and both the bar and the posterior process are exca-
vated by the renal depression. In the Odontophorus type the renal bar
is well developed and broad, and the bar, but not the posterior process,
is excavated by the renal depression.

THE DENDRORTYX GROUP
Dendrortyx. This genus appears to be the most primitive, as well as
the most aberrant genus of the New World quails.
Numerous characters of Dendrortyx are suggestive of those in less
advanced gallinaceous families as outlined by Holman (MS). The dor-
sal intermuscular line of the coracoid is sharply raised only along its
distal fifth. The ventral intermuscular line of the coracoid terminates
in the middle of the distal border of the sterno-coracoidal process.
The dorsal face of the coracoid has a round deep fossa below the
tubercle for the ligamentum sterno-coracoideum dorsale and above
the external end of the sternal facet. This is probably a remnant of
the deep pneumatic fossa found in this position in several less ad-
vanced families. The tip of the sterno-coracoidal process of the cora-
coid lacks a terminal knob. The depression on the medial side of the
pectineal process of the pelvis is perforated by a foramen. The fossa
II of the humerus is indistinct. The hypotarsus has only 1 closed cal-
caneal canal.
At the same time Dendrortyx is the most aberrant genus of the
Odontophoridae, for it has 30 unique characters excluding intermem-
bral proportions, and 41 unique characters including these propor-
tions. The distinctive intermembral proportions in Dendrortyx re-
flect the fact that the leg bones are long in comparison with the wing
bones, the ilium long in comparison with the sternum, and the height
of the sternal carina low in comparison with the length of the ilium.







Vol. 6


BULLETIN FLORIDA STATE MUSEUM










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HOLMAN: OSTEOLOGY OF QUAILS


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BULLETIN FLORIDA STATE MUSEUM


Dendrortyx has the largest measurements in the Dendrortyx group.
Philortyx. Philortyx is possibly the closest living relative of Dendror-
tyx, with which it shares the four following skeletal characters: The
capital groove of the humerus is deeply excavated and has its internal
margin strongly elevated. The height of the middle of the shaft of
the ulna is about 85 percent of the height of the shaft just distal to
the external cotyla. The pisiform process of the carpometacarpus is
round in internal view. The external condyle of the ulna has its ven-
tral border flattened. Philortyx has 8 unique characters but, aside
from its small size, it does not show striking skeletal modifications.

Oreortyx. This genus appears to show more relationships to Colinus,
Lophortyx, and especially to Callipepla (on characters of the scapula
and in pelvic proportions), than it does to Dendrortyx and Philortyx.
Oreortyx has 6 unique characters, but other than its relatively large
measurements, it shows no particularly noteworthy skeletal modifi-
cations.

Callipepla, Colinus, and Lophortyx. The genera of this complex often
cannot be separated on the basis of individual bones. Their skeletal
structure is so similar that Callipepla, Colinus, and Lophortyx must
have diverged relatively recently. Subgeneric rather than generic
designations might better reflect their taxonomic status, were it not for
their strongly differing external morphology.
Other studies support this view. Bailey (1928) reports a hybrid
Callipepla squamata X Lophortyx gambelii. Sibley (1960) states that
species of Colinus, Lophortyx, and Callipepla hybridize in various
combinations and reports that Callipepla, Colinus, and Lophortyx
share a characteristic egg-white protein profile different from that of
Oreortyx.

THE ODONTOPHORUS GROUP
Odontophorus. This genus appears to be the most primitive of the
Odontophorus group. Several important characters of Odontophorus
are found in less advanced gallinaceous families (Holman, MS). The
pneumatic fossa of the humerus has a perforated inner shelf that ex-
tends from the external bicipital surface to the medial bar in Odon-
tophorus gujanensis. Fossa II of the humerus is indistinct. The
hypotarsus has only one closed calcaneal canal in Odontophorus stel-
latus.
Some noteworthy characters occur on the pelvis of certain speci-
mens of Odontophorus. The posterior iliac crest of the pelvis has a


Vol. 6






HOLMAN: OSTEOLOGY OF QUAILS


prominence in all Odontophorus studied, with the exception of one
individual of Odontophorus gujanensis. A ridge originates on the pos-
terior iliac crest at the level of the ilio-ischiac fenestra, continues ven-
trad and posteriad toward the posterior border of the ilium, and has
a prominence that arises from its posterior surface in one individual of
Odontophorus gujanensis.
These characters have interesting implications for they are associ-
ated with the Dendrortyx-type pelvis and are not found in any other
genus in the Odontophorus group. Dendrortyx certainly appears to
be the most primitive genus of New World quails, and thus it seems
likely that the Dendrortyx-type pelvis evolved earlier than the Odon-
tophorus type. Therefore the occasional occurrence of these charac-
ters in a primitive genus of the Odontophorus group is not surprising.
Further substantiating the primitive nature of Odontophorus are
the three characters it shares with Dendrortyx only, namely: large size,
an indistinct fossa II of the humerus, the wing of the trochlea for digit
IV is depressed anteroposteriorly and the entire trochlea is longer than
deep.
Odontophorus is a rather generalized form with much specific
variation and with only 2 unique characters.

Dactylortyx. This genus has 10 unique characters but still does not
show as many skeletal modifications as the following two genera.
Perhaps the most important skeletal modifications in Dactylortyx are
the tendency toward the loss of the pectineal process with the com-
plete loss of the depression on the medial side of the pectineal process
and the elongation of the bones of the wing in comparison with the
bones of the leg. Dactylortyx is relatively large. It appears to be
more closely related to Cyrtonyx, somewhat more distantly related to
Rhynchortyx, and most distantly related to Odontophorus on the
basis of several qualitative characters as well as on intermembral pro-
portions.

Cyrtonyx. This form has 15 unique characters. Important skeletal
modifications include the strong deflection of the acromion process of
the scapula, the weak development of the intermetacarpal tubercle
of the carpometacarpus, the narrow angle of the ischium to the ventral
edge of the posterior iliac crest, and, as in Dactylortyx, the elongation
of the bones of the wing in comparison to the bones of the leg. Cyr-
tonyx also is relatively large. It appears most closely related to Dac-
tylortyx, somewhat more distantly related to Rhynchortyx, and most
distantly to Odontophorus.






BULLETIN FLORIDA STATE MUSEUM


Rhynchortyx. This most aberrant genus of the Odontophorus group
has 25 unique characters. Important skeletal modifications include
the short, deep, rostrum and the development of a third distinct cal-
caneal ridge on the hypotarsus. Rhynchortyx has the smallest measure-
ments of any genus in the Odontophorus group. In several qualitative
characters, as well as on intermembral proportions, it appears more
closely related to Cyrtonyx and Dactylortyx than to Odontophorus.

REMARKS
In summary Dendrortyx appears to be at once the most primitive and
aberrant genus of the Odontophoridae. Philortyx is possibly its closest
relative. Oreortyx is closely related to Colinus, Lophortyx, and espe-
cially to Callipepla. Callipepla, Colinus, and Lophortyx are so similar
that on osteological grounds alone they could be merged in a single
genus.
Odontophorus appears to be the most primitive genus in the Odon-
tophorus group. Dactylortyx and Cyrtonyx show several similarities
and appear closer to Rhynchortyx than to Odontophorus. Rhynchor-
tyx is the most aberrant genus of the Odontophorus group.
On the basis of the present study, the following rearrangement is
proposed:


Sequence of genera
in Peters (1934)
Dendrortyx
Oreortyx
Callipepla
Lophortyx
Philortyx
Colinus
Odontophorus
Dactylortyx
Cyrtonyx
Rhynchortyx


Proposed sequence of genera
Group 1
Dendrortyx
Philortyx
Oreortyx
Callipepla
Colinus
Lophortyx
Group 2
Odontophorus
Dactylortyx
Cyrtonyx
Rhynchortyx


Vol. 6





HOLMAN: OSTEOLOGY OF QUAILS


FOSSIL RECORD OF NEW WORLD QUAILS
American quails are reported from Oligocene to Recent times, but
the single Oligocene form is indeterminate at the generic and specific
levels. Modern genera are first recorded from the Miocene, and
modern species first appear in the Pleistocene.

TERTIARY FORMS
ODONTOPHORIDAE, INDETERMINATE
HORIZON AND LOCALITY. Oligocene: silts of Orellan age in the
Cedar Creek faces of the White River formation. From SWY4 sec.
12, T. 11 N., R. 54 W., Logan County, Colorado (Tordoff, 1951b).
MATERIAL. Distal end of a tarsometatarsus (Univ. Kansas Nat.
His. Mus.).
REMARKS. Tordoff (1951b) states that though the Oligocene fossil
is smaller than Colinus and Lophortyx, it most closely resembles these
genera. Some American genera of quails (and even some gallinaceous
families) are difficult to differentiate on the basis of the distal portion
of the tarsometatarsus, and the reported affinities of the Oligocene
fragment should be regarded with some caution.

Miortyx A. H. Miller, 1944
The genus is known from a single species recorded from the Lower
Miocene of South Dakota.
Miortyx teres A. H. MILLER, 1944
HORIZON AND LOCALITY. Lower Miocene: late Arikareean stage of
Rosebud formation. From Flint Hill quarry, Bennett County, South
Dakota (A. H. Miller, 1944).
MATERIAL. Type, proximal three-fourths of right humerus (Univ.
California Mus. Paleont. 34453). Referred specimen, distal end of left
tibiotarsus.
DIAGNOSIS. "Proximal end of humerus similar to that of Oreortyx
but decidedly larger; capital groove sharply defined and narrower,
especially at margin of head; contour of head more smoothly rounded,
less indented by capital groove; ligamental furrow on palmar surface
deeper; tubercle on middle of deltoid crest proportionately larger."
MEASUREMENTS. "Type, greatest width of head of humerus per-
pendicular to axis of upper part of shaft, 12.9 mm.; least width of
shaft, in mediolateral direction, 5.0."
REMARKS. A. H. Miller (1944) points out: "The most distinctive
feature of Miortyx is the rounded contour of the head with consequent






BULLETIN FLORIDA STATE MUSEUM


shallow depression between the articular surface of the head and the
internal tuberosity. Dendrortyx of Central America approaches, but
by no means equals Miortyx with respect to lesser indentation of the
head contour by the capital groove."
In the living Odontophoridae fossa II of the humerus is usually
well developed. Dendrortyx and Odontophorus are apparently less
specialized exceptions in which fossa II is obsolete. The drawing of
the type specimen of Miortyx teres (A. H. Miller, 1944: 94, fig. 7) in-
dicates that fossa II, though not obsolete, is rather weakly developed.

Cyrtonyx Gould, 1844
The genus is reported from the upper Miocene of California and Ne-
braska, and from the Pleistocene of Aramberri, Nuevo Leon, Mexico.
Cyrtonyx cooki WETMORE, 1934
HORIZON AND LOCALITY. Upper Miocene: upper Sheep Creek beds.
From 17 miles south of Agate, Sioux County, Nebraska (Wetmore,
1934).
MATERIAL. Type, distal half of left humerus (collection of Harold
J. Cook, HC 647).
DIAGNOSIS. "Distal end of humerus similar to that of Cyrtonyx
montezumae mearnsi Nelson, but about one-fourth larger; ectepicon-
dyle relatively reduced."
MEASUREMENTS. Greatest transverse breadth across condyles 9.5
mm.; least transverse breadth of shaft 4.6 mm.
REMARKS. Wetmore (1934) reports that the fossil is a northward
extension of Cyrtonyx, which occurs today from central Arizona and
central Texas southward into Guatamala.
Cyrtonyx tedfordi L. MILLER, 1952
HORIZON AND LOCALITY. Upper Miocene: Barstow formation, Lake
bed horizon. From near middle of SE4 NW/4 sec. 15, T. 11 N., R. 2
W., near the town of Barstow, California (L. Miller, 1952).
MATERIAL. Type, right carpometacarpus (Univ. California Mus.
Paleont. 42223).
DIAGNOSIS. "Very similar to male of Cyrtonyx montezumae mearn-
si, but much stouter, and with intermetacarpal tubercle much less de-
veloped."
REMARKS. This fossil needs restudy. The photograph of the type
carpometacarpus (L. Miller, 1952: 299, fig. 2) shows the following
characters: Pisiform process reduced, and at the level of its ligamental
attachment. Intermetacarpal tubercle obsolete, represented only by


Vol. 6





HOLMAN: OSTEOLOGY OF QUAILS


a minute point. Carpal trochlea with its external rim continuing dis-
tad beyond ligamental notch. These are strong characters of the
family Cracidae and do not occur in any living genus of American
quails. Possibly this fossil may represent a small cracid.

Colinus Goldfuss, 1820
The genus is reported from the late Pliocene of Arizona and Kansas,
the Pleistocene of Florida and Texas, and prehistoric deposits from
caves in Yucatan, Florida, Texas, and Tennessee.
Colinus sP.
HORIZON AND LOCALITY. Upper Pliocene. From sec. 22, T. 17 S.,
R. 20 E., about two miles south of Benson, Arizona (Wetmore, 1924).
MATERIAL. Distal portion of tarsometatarsus (U. S. Natl. Mus.).
REMARKS. The fossil has a smaller and narrower middle trochlea
than Callipepla and a lower distal foramen than either Colinus virgini-
anus or Lophortyx gambelii (Wetmore, 1924).
Colinus hibbardi WETMORE, 1944
HORIZON AND LOCALITY. Upper Pliocene: Rexroad formation.
From Locality 3, Rexroad fauna, Meade County, Kansas (Wetmore,
1944).
MATERIAL. Type, distal portion of right tarsometatarsus, with
most of the outer trochlea missing (Univ. Kansas Mus. Vert. Paleont.
3981). Referred specimens, distal end of right humerus and distal end
of badly worn left humerus.
DIAGNOSIS. "Distal end of tarsometatarsus (figs. 4 and 5) similar to
that of modern Colinus virginianus (Linnaeus) but decidedly larger;
shaft stronger, and more heavily lined by the tendinal grooves."
MEASUREMENTS. Type: "Transverse breadth of shaft below cen-
ter, 3.1 mm.; transverse breadth across trochlea (approximate), 7
mm.; transverse breadth of middle trochlea, 2.9 mm."
REMARKS. Wetmore (1944) reports that the type tarsometatarsus
shows more angular development of the posterior side of the middle
trochlea than in Lophortyx, stouter form throughout than in Callipepla,
and larger trochlea than in Cyrtonyx. The referred humerus (Univ.
Kansas Mus. Vert. Paleont. 3997) shows similarities to Colinus virgini-
anus, but the brachial depression is larger, and the ridge that borders
it is longer and extends farther up the shaft. In size it is similar to
adult male Cyrtonyx montezumae mearnsi, but the trochlea are defi-
nitely smaller. The radial trochlea is larger than in Callipepla. All of
the above fossils are larger than Colinus virginianus.





BULLETIN FLORIDA STATE MUSEUM


Tordoff (1951a) reports on 32 bones of Colinus hibbardi from the
type locality. Fossil elements include 5 vertebrae, 3 sterna, 7 cora-
coids, 1 humerus, 5 ulnae, 7 carpometacarpi, and 4 tarsometatarsi.
Tordoff finds that the fossils are similar to the bones of Recent Colinus
virginianus but larger, and the wing bones are longer in relation to
the leg bones. Subjective differences include the following characters:
Ventral intermuscular line of coracoid less prominent and less sharply
defined than in Recent C. virginianus. Shaft of humerus straighter
and heavier, and brachial depression larger. Impression of M. brachi-
alis anticus on ulna deeper and more sharply outlined. Intercotylar
prominence of tarsometatarsus more nearly round, depression just
over posterior edge of internal cotyla more angular and sharply
marked, and the ridge leading proximad from the outer proximal fora-
men more sharply defined.

ODONTOPHORIDAE, INDETERMINATE
HORIZON AND LOCALITY. Upper Pliocene. From sec. 25, T. 18
S., R. 21 E., about 14 miles southeast of Benson, Arizona (Wetmore,
1924).
MATERIAL. Proximal end of humerus (U. S. Natl. Mus.).
REMARKS. Wetmore notes that this fragment is large as in Oreor-
tyx, but has characters resembling Colinus. It seems possible that the
specimen represents C. hibbardi.

Lophortyx Bonaparte, 1838
The genus is reported from the middle Pliocene of Oregon, the Pleisto-
cene of California, and from archaeological sites in California and
Arizona.

Lophortyx shotwelli BRODKORB, 1958
HORIZON AND LOCALITY. Middle Pliocene: Hemphillian stage.
From tuffaceous sandstone on the east bank of McKay Reservoir, about
five miles south of Pendleton, Umatilla County, Oregon (Brodkorb,
1958).
MATERIAL. Type, proximal portion of left humerus (Univ. Oregon
Mus. Nat. Hist. F-3611). Referred material, proximal portion of left
humerus, and distal portion of two left humeri.
REMARKS. Through the kindness of the University of Oregon Mu-
seum of Natural History the type material was made available for
study. A description based on this re-examination is as follows:
Humerus with 1) greatest proximal width 9.8 mm. in type specimen (re-


Vol. 6





HOLMAN: OSTEOLOGY OF QUAILS


ferred specimen broken). 2) Greatest distal width of humerus 7.3 mm.
3) Pneumatic fossa moderately elongate, elliptical, reduced in its distal
extent, and with a vestigial, perforated, inner shelf. 4) Median crest
strongly developed, elevated throughout. 5) Capital groove moder-
ately excavated with its internal margin moderately elevated. 6) Mar-
gin between head and internal tuberosity moderately concave. 7)
Head about as long as broad in anconal view. 8) Ridge along medial
border of fossa II moderately swollen, elongate, and with its apex
acute. 9) Fossa II well developed. 10) Latissimus ridge originating
on lateral edge of shaft, but swinging in onto anconal face proximally.
11) External tuberosity weakly inflated. 12) Bicipital crest arising
gently from shaft, narrowly rounded, with its distal border lacking a
groove. 13) Deltoid crest moderately developed, slightly inflected,
and with its summit flattened in type specimen (knoblike in referred
specimen). 14) Entepicondyle at level of internal condyle. 15) En-
tepicondylar prominence moderately produced, with its pit placed
on internal face. 16) Depression of M. brachialis anticus moderately
excavated.
Several of the above characters are noteworthy. Perhaps most in-
teresting is the vestigial, perforated, inner shelf of the pneumatic fossa
of the humerus. A complete imperforate inner shelf is present in most
other families of the Galliformes, but has been lost in all living New
World quails with the exception of Odontophorus gujanensis, which re-
tains a perforated inner shelf. Lophortyx shotwelli differs from all
Recent species of Lophortyx in characters 3, 4, 11, and 14 above, and
shares character 7 with Recent Dendrortyx, Philortyx, and Lophortyx
douglasii only.
PLEISTOCENE FORMS
Dendrortyx Gould, 1844
The genus is reported from the Pleistocene of Nuevo Leon, Mexico,
and tentatively from a prehistoric deposit in a Yucatan cave (Fisher,
1953).

?Dendrortyx
HoRIZON AND LOCALITY. Pleistocene. From San Josecito Cavern,
province of Aramberri, state of Nuevo Ledn, Mexico (L. Miller, 1943).
MATERIAL. A single femur (Univ. California Mus. Paleont.).
REMARKS. L. H. Miller reports this femur as larger and heavier
than that of a Recent specimen of Dendrortyx leucophrys nicaraguae,
but refers it only tentatively to this genus.





BULLETIN FLORIDA STATE MUSEUM


Oreortyx Baird, 1858
Reported from the late Pleistocene of California, and from the Quat-
ernary (probably prehistoric) of a New Mexico cave.

Oreortyx picta (DOUGLAS, 1829)
HORIZON AND LOCALITY. (a) Late Pleistocene. From Potter Creek
Cave, Shasta County, California (L. Miller, 1911). (b) Late Pleisto-
cene. From Samwel Cave, Shasta County, California (L. Miller,
1912). (c) Late Pleistocene. From Hawver Cave, Eldorado County,
California (L. Miller, 1911).
MATERIAL. (a) Potter Creek Cave; 2 bones. (b) Samwel Cave;
not specified. (c) Hawver Cave; 14 bones. Bones from the above lo-
calities are at the University of California Museum of Paleontology.

Neortyx, new genus
Figure 3
TYPE OF GENUS. Neortyx peninsularis, new species.
DIAGNOSIs. Differs from living genera of New World quails as
follows. 1) Femur with proximal end narrow throughout, with head
and trochanter small, but with shaft expanding abruptly into enlarged
distal end (living genera have proximal end wider throughout, with
head and trochanter larger, and with shaft expanding less abruptly
into less enlarged distal end). 2) Anterior border of external condyle
extending well onto distal portion of shaft as a strong ridge (anterior
border of external condyle ends more abruptly on distal portion of
shaft as a weak ridge in living genera). 3) Internal lip of external
condyle reflected (deflected in living genera). 4) Dorsolateral border
of iliac facet short (longer in living genera).
Closest to more specialized genera of the Dendrortyx group in
having 1) head neither bent dorsad nor rotated posteriad, thus differ-
ing from Dendrortyx and Rhynchortyx. 2) Neck moderately long, thus
differing from Dendrortyx. 3) Iliac facet without compression, with
posterior border convex, thus differing from Dendrortyx. 4) Dorsal
crest of trochanter broken. 5) Trochanteric ridge 24 percent of length
of shaft, thus differing from Rhynchortyx and Dactylortyx. 6) Distal
depth 83 percent of distal width, thus differing from Philortyx and
Rhynchortyx. 7) Internal condyle moderately wide, and with its an-
terior border much produced throughout, thus differing from Philor-
tyx, Cyrtonyx, and Dactylortyx. 8) External condyle compressed and
rounded anteriorly, thus differing from Dendrortyx.


Vol. 6






HOLMAN: OSTEOLOGY OF QUAILS


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BULLETIN FLORIDA STATE MUSEUM


Neortyx peninsularis, NEW SPECIES
HOLOTYPE. Left femur, complete except for broken dorsal crest
of trochanter (Brodkorb collection [PB 1224], Univ. Florida). Pleisto-
cene: Arredondo clay equivalent, in pit of Dixie Lime Products Com-
pany in the SW corner NW4 sec. 14, T. 13 S., R. 21 E., Marion County,
Florida, 1 mile southeast of the town of Reddick. Collected by Pierce
Brodkorb 9 August 1954.
MEASUREMENTS OF HOLOTYPE. Length through articular facet, 45.8
mm., greatest proximal width, 7.5 mm., distal width through condyles,
7.6 mm., greatest proximal width/length 16.4 percent, distal width
through condyles/greatest proximal width 101.3 percent, proximal
width of shaft/width of shaft just proximad to internal condyle 65.2
percent.
DIAGNOSIS. Femur larger than in Philortyx fasciatus, Oreortyx
picta, Callipepla squamata, Colinus suilium, C. virginianus, C. nigro-
gularis, C. leucopogon, Lophortyx californica, L. gambelii, L. douglasii,
Dactylortyx thoracicus, and Cyrtonyx montezumae; smaller than in
Dendrortyx leucophrys, Odontophorus gujanensis, 0. stellatus, and
0. guttatus (table 1).
REFERRED MATERIAL. The following elements from the Pleisto-
cent of Reddick, Florida, are referred to the above species. One com-
plete and 2 fragmentary right coracoids (PB 478, 1275, and 1974), and
2 fragmentary right ulnae (PB 935 and 1229).
Coracoid.-Internal distal angle short, rounded terminally, and
with its dorsal surface flattened (internal distal angle elongate, pointed
terminally, and with its dorsal surface round in living genera). Length
29.0 mm., proximal width 4.3 mm., distal width (two specimens), 8.7-
8.8 mm., distal width/length 30.3 percent, proximal width/distal
width 48.9 percent, proximal width/length 14.8 percent.
Ulna.-Olecranon process only moderately produced (strongly pro-
duced in most living genera, weakly produced in Rhynchortyx).
Greatest proximal width 6.0-6.3 mm.
REMARKS. This is the second extinct genus of New World quails
described and brings the total of extinct and living genera to 12. The
type specimen shows several resemblances to Oreortyx, Callipepla,
Colinus, and Lophortyx, but is quite distinct from these forms, as it
has four unique characters. The referred elements show two unique
characters, but otherwise are similar to birds in the Callipepla-Colinus-
Lophortyx complex in qualitative characters as well as in size.


Vol. 6






HOLMAN: OSTEOLOGY OF QUAILS


Colinus Goldfuss, 1820
Colinus suilium BRODKORB, 1959
HORIZON AND LOCALITY. Pleistocene: Arredondo clay member of
Wicomico formation. From Pits I and II in limestone quarry in NWV4
sec. 22, T. 10 S., R. 9 E., near Arredondo, Alachua County, Florida
(Brodkorb, 1959).
MATERIAL. Holotype, Pit I: complete left humerus, (Brodkorb
collection, PB 1291). Paratype, Pit I: complete right humerus (PB
757). Referred material, Pit I: 1 sternum, 2 coracoids, 4 humeri, 3
ulnae, 1 carpometacarpus, 1 synsacrum, 3 tibiotarsi, 3 tarsometatarsi.
Pit II: 2 coracoids, 1 scapula, 1 ulna, 4 humeri, 2 femora, 3 tibiotarsi,
and 1 tarsometatarsus (Brodkorb collection).
DIAGNOSIS. For the original description see Brodkorb (1959: 276).
STATUS OF THE ARREDONDO CLAY. The Arredondo clay member of
the Wicomico formation consists mainly of blue clay streaked with
yellow that weathers to brown. It was deposited under terrestrial
conditions of sedimentation and contains fossils of freshwater gastro-
pods and vertebrates. Arredondo clay lies beneath the Wicomico ter-
race, a reddish-brown marine sand. The exact age of the Wicomico
formation is uncertain, but possibly it represents either Yarmouth or
Sangamon interglacial deposition (Brodkorb, personal communica-
tion). Therefore, the Arredondo clay may represent either Kansas or
Illinoian times.
OTHER LOCALITIES. Fossil bones assigned to the species Colinus
suilium from other fossil localities in Florida listed in the next para-
graphs are discussed in a later section on variation in fossil Colinus:
(a) Williston, Florida.-Pleistocene: Arredondo clay. From sink-
hole in Connell and Shultz Limerock Company mine, 0.9 miles north
of Seaboard Airline Railroad, Williston, Levy County, Florida (Hol-
man, 1959a and 1959c): 1 rostrum, 1 coracoid, 5 humeri, 1 ulna, 2
carpometacarpi, 4 femora, 3 tibiotarsi, and 1 tarsometatarsus (Brod-
korb collection).
(b) Orange Lake, Florida.-Pleistocene: Arredondo clay and Arre-
dondo clay-Wicomico sand contact. From sinkhole in limestone quar-
ry near center of sec. 33, T. 12 S., R. 21 E., Marion County, Florida,
2 miles south of the town of Orange Lake (Holman, 1959b). Arre-
dondo clay: 8 coracoids, 4 scapulae, 9 humeri, 2 ulnae, 1 carpometa-
carpus, 10 femora, 5 tibiotarsi, 5 tarsometatarsi. Arredondo clay-
Wicomico sand contact: 1 femur (Brodkorb collection).
(c) Reddick, Florida.-Pleistocene: Arredondo clay equivalent.
From localities A, B, C, and undesignated exact localities (either A or







BULLETIN FLORIDA STATE MUSEUM


B) in pit of Dixie Lime Products Company in the SW corner NW/4
sec. 14, T. 13 S., R. 21 E., Marion County, Florida, 1 mile southeast
of the town of Reddick (Brodkorb, 1957).
Locality A: 1 sternum, 5 coracoids, 9 humeri, 2 ulnae, 2 femora,
5 tibiotarsi, 2 tarsometatarsi. Locality B: 3 coracoids, 6 humeri, 7
femora, 2 tibiotarsi, 7 tarsometatarsi. Locality C: 1 rostrum, 2 cora-
coids, 15 humeri, 4 ulnae, 9 carpometacarpi, 1 synsacrum, 8 femora,
4 tibiotarsi, 4 tarsometatarsi. Locality A or B undesignatedd): 1 ros-
trum, 3 sterna, 68 coracoids, 9 scapulae,. 138 humeri, 63 ulnae, 30 carpo-
metacarpi, 5 synsacra, 90 femora, 77 tibiotarsi, 61 tarsometatarsi (Brod-
korb collection).
The total of 644 bones from Reddick is relatively a tremendous
number of individual fossil bird bones. The 74 left humeri from Red-
dick show that a minimum of 74 individuals is represented.
(d) Haile II, Florida.-Pleistocene: Arredondo clay. From locality
II B in NW/4 sec. 25, T. 9 S., R. 13 E., Alachua County, Florida, near
the village of Haile (Auffenberg, 1955). A fragmental ulna and tibio-
tarsus (collection of Howard Hutchison, Univ. Florida).
(e) Oakhurst Quarry, Florida.-Pleistocene. From Oakhurst Lime-
stone Quarry, 2 miles southeast of Ocala Post Office, Marion County,
Florida. A single left humerus (Brodkorb collection). As no other
extinct vertebrates have been taken from this locality its age is un-
certain.
(f) Zuber, Florida.-Pleistocene: Arredondo clay equivalent. Dixie
Lime Products Company mine, Zuber, Marion County, Florida. One
femur (Brodkorb collection).
(g) Eichelberger Cave, Florida.-Pleistocene (stratigraphy unclear).
From Locality A in solution cave in NW corner of sec. 2, T. 17 S.,
R. 22 E., Marion County, Florida, 2 miles southwest of the town of
Belleview (Brodkorb, 1956): 1 humerus, 1 femur, 1 tibiotarsus, and 1
tarsometatarsus (Brodkorb collection).
(h) Lake Monroe, Florida.-Pleistocene. From Locality N, dredged
from mouth of Lake Monroe, Volusia County, Florida, associated with
extinct Pleistocene vertebrates: 1 humerus.
Specimens from localities (d) through (h) are referred tentatively
to C. suilium as they show an approach to C. virginianus in some re-
spects.

Colinus virginianus (LINNAEUS, 1758)
Fossil bones assigned to the species Colinus virginianus listed in the
next paragraphs are discussed in a later section on variation in fossil
Colinus.


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HOLMAN: OSTEOLOGY OF QUAILS


(a) Kanapaha, Florida.-Pleistocene (stratigraphy unclear). From
limestone quarry in sec. 22, R. 19 E., T. 10 S., Alachua County, Florida,
about 1.5 miles SW of the village of Arredondo (Brodkorb, 1959): 2
humeri, 1 carpometacarpus, 1 femur, and 1 tibiotarsus (Brodkorb col-
lection). The bones are from tan sand in solution holes in the Ocala
limestone, but the relationship of the deposit to the Wicomico terrace
is not well marked.
(b) Sabertooth Cave, Florida.-Pleistocene: post-Wicomico. From
solution cave in NW/4 SWY4 sec. 33, T. 18 S., R. 18 E., Citrus County,
Florida, 1.2 miles NW of the town of Lecanto: 2 humeri, 4 carpometa-
carpi, 3 femora, 2 tibiotarsi (Brodkorb collection); and 2 humeri, 2
femora, and 1 tarsometatarsus (Holmes collection, see Wetmore, 1931).
Holman (1958) reports incorrectly that the fossils lie in matrix derived
from the Pamlico terrace. The matrix was derived from the Wicomico
terrace.
(c) Seminole Field, Florida.-Pleistocene: post-Wicomico. From
Joes Creek, 46th Avenue N., 71st St. N., St. Petersburg, Pinellas
County, Florida (Wetmore, 1931): 6 humeri and 1 ulna (Holmes col-
lection, see Wetmore, 1931).
(d) Melbourne, Florida.-Pleistocene: post-Wicomico. From line
of contact between strata 1 and 2, Country Club Golf Links, near
Melbourne, Brevard County, Florida (Wetmore, 1931). Two humeri
collected by J. W. Gidley in 1928 appear to be lost at present.
(e) Vero, Florida.-A heterochronic locality: stratum 2 (with extinct
vertebrates), Pleistocene, pre-Pamlico; stratum 3 (without extinct verte-
brates), Pleistocene or Recent, post-Pamlico; "West of Bridge," age
uncertain. All sites in the center of sec. 35, T. 32 S., R. 39 E., Vero
Beach, Indian River County, Florida (Weigel, MS). Stratum 2: 2
humeri, 1 coracoid, and 1 tibiotarsus; stratum 3: 2 humeri, 1 ulna, and
1 tibiotarsus; "West of Bridge": 1 humerus (Brodkorb collection).
(f) Warren's Cave, Florida.-?Prehistoric. From floor of Warren's
Cave, 8.5 miles northwest of Gainesville, Alachua County, Florida:
1 scapula, 1 coracoid, and 1 tarsometatarsus (Brodkorb collection).
(g) Friesenhahn Cave, Texas.-Late Pleistocene. From northern
Bexar County, Texas (Milstead, 1956): 4 humeri, 1 ulna, and 2 tibio-
tarsi (Texas Mem. Mus. collection 933-428, 933-447, 933-2186, 933-3042,
933-3199, 933-3041, and 933-3531).

Lophortyx Bonaparte, 1838
Lophortyx californica (SHAW, 1798)
HORIZON AND LOCALITY. (a) Pleistocene. From Rancho La Brea
asphalt beds in Los Angeles, Los Angeles County, California (Howard,






BULLETIN FLORIDA STATE MUSEUM


1930; Miller and DeMay, 1942). (b) Pleistocene. From Fauna I and
Fauna II of asphalt beds, McKittrick, California (L. Miller, 1935). (c)
Pleistocene. From asphalt beds of Carpinteria, California (DeMay,
1941). (d) Pleistocene. San Pedro and Palos Verdes formations.
San Pedro, California (L. Miller, 1914 and 1930; Howard, 1944 and
1949). (e) Pleistocene. From Hawver Cave, Eldorado County, Cali-
fornia (L. Miller, 1911).
MATERIAL. (a) Rancho La Brea: 101 individuals. (b) McKittrick,
Fauna I: 3 bones; Fauna II: 93 bones. (c) Carpinteria: 9 individuals,
102 bones. (d) San Pedro, San Pedro formation: 1 coracoid, 2 humeri,
1 tibiotarsus; Palos Verdes formation: 6 bones. (e) Hawver Cave: 17
bones. Fossils from all the above localities are in the University of
California Museum of Paleontology. Fossils from localities (a) and
(d) are also in the Los Angeles County Museum. Lophortyx cali-
fornica is the most plentiful fossil quail in the western United States.

Cyrtonyx Could, 1844
Cyrtonyx montezumae (VIGORS, 1830)
HoRIzON AND LOCALITY. Pleistocene. From San Josecito Cavern,
province of Aramberri, Nuevo Leon, Mexico (L. Miller, 1943).
MATERIAL. Eighty bones, including limb bones, a coracoid, and
two crania (Univ. California Mus. Paleontol.)
REMARKS. According to L. Miller (1943) the fossil sample is close
to the subspecies C. montezumae mearnsi.

Odontophorus Vieillot, 1816
The genus is reported from the Pleistocene of various caves in Brazil,
and from a prehistoric site in a Yucatan cave.

Odontophorus gujanensis (GMELIN, 1789)
HORIZON AND LOCALITY. Pleistocene. From four cave deposits
(Lapa da Escrivania, Lapa da Marinho, Lapa do Periperi, and Lapa
Vermelha) near Lagoa Santa, Minas Geraes, Brazil (Winge, 1887).
MATERIAL. A few bones from each cave (Copenhagen Mus.)

FORMS FROM ARCHAEOLOGICAL SITES
Remains of species of American quails living today are known from
several archaeological sites. Dendrortyx leucophrys is listed question-
ably from a cave in Yucatan (Fisher, 1953). Oreortyx picta is reported
from a cave in New Mexico (Wetmore, 1932b). Colinus virginianus is
identified from caves in Tennessee and Texas (Shufeldt, 1897; Wet-


Vol. 6





HOLMAN: OSTEOLOGY OF QUAILS


more, 1933), and C. nigrogularis is reported from a cave in Yucatan
(Fisher, 1953). Lophortyx californica and L. gambelii are listed from
Indian sites in California and Arizona (DeMay, 1942; Wetmore, 1932a).
Odontophorus guttatus is identified questionably from a cave in Yuca-
tan (Fisher, 1953).

VARIATION IN FOSSIL Colinus
Wetmore (1944) named Colinus hibbardi from the upper Pliocene of
Meade County, Kansas, and Brodkorb (1959) described Colinus su-
ilium from the pre-Wicomico Pleistocene of Arredondo, Florida. These
species were described on the basis of large size and several qualita-
tive characters. Additional specimens of C. hibbardi and C. suilium
were reported by Tordoff (1951a) and Holman (1959c). As Brodkorb
(1960) points out, in all probability these forms are the temporal equiv-
alents of living Colinus virginianus, and we are dealing with a con-
tinuum. Because the living species of Colinus are almost identical to
each other in osteological characters and exhibit subspecific variation
in size, it is extremely difficult to decide where to draw the taxonomic
line between certain fossil populations of Colinus. It should be
pointed out that the samples of fossil bones from many localities are
too small to allow statistically significant analysis.
QUALITATIVE CHARACTERS
CORAcom. The head of the coracoid is reflected in the single
fossil of C. hibbardi, in 63 percent of 62 fossils of C. suilium from Ar-
redondo, Orange Lake, and Reddick, in 92 percent of 86 modern speci-
mens of C. virginianus and in the 2 specimens of modern C. nigrogu-
laris. It is inflected in 2 of 3 specimens of modern C. leucopogon.
The procoracoid process is weakly developed in C. hibbardi and is
stronger in C. suilium, C. virginianus, C. nigrogularis, and C. leucopo-
gon.
SCAPULA. The dorsal depression just media to the glenoid facet
is deep in 67 percent of 15 fossils of C. suilium from Arredondo, Orange
Lake, and Reddick, but is shallow in 77 percent of 81 modern speci-
mens of C. virginianus. The depression is deep in 1 of 2 specimens of
living C. nigrogularis, and in 2 of 3 specimens of living C. leucopogon.
The bridge between the acromion process and glenoid facet is
usually about one-half the width of the glenoid facet in C. suilium, C.
virginianus, and C. leucopogon, but is less than one-third the width of
the glenoid facet in C. nigrogularis.
The acromion process is strongly inflected in 62 percent of 13 fos-
sils of C. suilium from Arredondo, Orange Lake, and Reddick. In the





BULLETIN FLORIDA STATE MUSEUM


living species the process is only slightly inflected in 72 percent of 80
specimens of C. virginianus, in 1 of 2 C. nigrogularis, and in 2 of 3
C. leucopogon.
HUMERUS. The ridge along the medial border of fossa II is mod-
erately swollen in fossil C. suilium, modern C. virginianus, and modern
C. leucopogon, but is only slightly swollen in modern C. nigrogularis.
The bicipital crest is pointed in 51 percent of 97 fossils of C. suilium
from Arredondo, Williston, Orange Lake, and Reddick. It is rounded
in 69 percent of 83 specimens of living C. virginianus, in the single
specimen of C. nigrogularis, and in the 3 specimens of living C. leu-
copogon.
The entepicondyle arises abruptly from the shaft in 20 percent
of 107 fossils of C. suilium from Arredondo, Williston, Orange Lake,
and Reddick. The entepicondyle arises gently from the shaft in 97
percent of 77 specimens of modern C. virginianus and in the 3 speci-
mens of C. leucopogon.
The depression for M. brachialis anticus is large in C. hibbardi,
and in 2 of 4 fossils of C. suilium from Arredondo, but it is small in
the living species of Colinus.
CARPOMETACARPUS. The process of metacarpal I is pointed in the
single fossil of C. hibbardi, and in 63 percent of 35 fossils of C. suilium
from Arredondo, Williston, Orange Lake, and Reddick. It is pointed
in 93 percent of 44 specimens of living C. virginianus, but is rounded
in the 3 specimens of C. leucopogon.
The pisiform process is much reflected dorsad and thus produced
well above the dorsal rim of the inner carpal trochlea in 25 percent
of 32 fossils of C. suilium from Arredondo, Orange Lake, and Reddick.
The process is slightly reflected dorsad and thus below or at the level
of the inner carpal trochlea in all living species.
The notch between the proximal border of the process of meta-
carpal I is deep in the single fossil of C. hibbardi and in 80 percent
of 35 fossils of C. suilium from Arredondo, Williston, Orange Lake,
and Reddick. The notch is slight in 71 percent of 48 specimens of
living C. virginianus and in 2 of 3 specimens of living C. leucopogon.
TIBIOTARSUs. The posterior margin between the external and in-
ternal articular surface is deeply notched in 28 percent of 18 fossils of
C. suilium from Arredondo, Orange Lake,.and Reddick. The notch
is shallow in all specimens of the 3 living species.

QUANTITATIVE CHARACTERS
Two types of size trends occur in the genus Colinus. A geographic
decrease in size occurs from north to south, with Michigan quail the


Vol. 6






HOLMAN: OSTEOLOGY OF QUAILS


largest, and southern Florida quail the smallest (table 2). A temporal
decrease in mean size occurs from late Pliocene to the present.
Colinus hibbardi from the upper Pliocene of Kansas is the largest
known Colinus, and quail from the Pleistocene of Florida are usually
larger than their Recent geographic representatives. Fossils assigned
to the species C. suilium from Arredondo, Williston, Orange Lake, and
Reddick, Florida, are decidedly larger than quail living in those places
today (table 8). They average as large or larger than Recent Colinus
virginianus from Michigan in most measurements (Arredondo, 19 of
20 measurements; Williston, 11 of 12; Orange Lake, 17 of 17; and
Reddick, 16 of 20). A few bones tentatively assigned to C. suilium
from Haile II, Oakhurst Quarry, Eichelberger Cave, and Lake Mon-
roe, Florida, are also similar in size to living C. v. virginianus from
Michigan, and a single bone from Zuber, Florida, tentatively assigned
to C. suilium, is similar in size to living C. v. virginianus from Illinois.
Fossils assigned to C. virginianus from Kanapaha, Saber-tooth
Cave, the Vero strata, and Warren's Cave, Florida, are about the size
of their Recent geographic representatives. Bones assigned to C. vir-
ginianus from the Pleistocene of Friesenhahn Cave, Texas, are only
slightly larger than Recent bobwhites from northern Florida.
Recent and fossil species of Colinus are inseparable on intermem-
bral and intramembral ratios with the exception of one fossil humerus
from Oakhurst Quarry near Ocala, Florida, which is slightly shorter
and stouter than in other Recent and fossil Colinus.

CONCLUSIONS
The American quails form a natural group that has undergone an in-
dependent evolution in the New World (Holman, MS). The following
hypothetical outline of the course of events is based on the osteology
of the living species and on the fossil record.
The families of living Galliformes were differentiated early in the
Tertiary period. Fossils representing the Cracidae (Gallinuloides East-
man; see Tordoff and Macdonald, 1957), the Phasianidae (Palaeortyx
Milne-Edwards), and the Tetraonidae (Palaeophasianus Shufeldt) are
recorded from the Eocene. The Meleagrididae (Meleagris antiqua
Marsh), and the Odontophoridae (indeterminate odontophorid, Tor-
doff, 1951b) are represented by Oligocene fossils.
The hypothetical ancestor of the Odontophoridae must have been
close to the stock from which the pheasants and grouse arose. It
probably was in this stage of skeletal evolution: The size was relatively
large, and the rostrum was short and deep. The sternum was charac-





BULLETIN FLORIDA STATE MUSEUM


terized by deep notches, long, parallel, anterior lateral processes, mod-
erately pneumatic sternal plate, and reduced dorsal foramen of the
manubrial spine. The coracoid had a ventrally overhanging brachial
tuberosity, with the dorsal intermuscular line slightly raised distally,
and the ventral intermuscular line encroaching on the sterno-cora-
coidal process, whose tip was without a terminal knob. The scapular
blade was elongate and had the beginning of a dorsal groove. The
humerus had a moderately large pneumatic fossa with a reduced inner
shelf, and fossa II was still indistinct. The carpometacarpus had a
moderately produced pisiform process and a well-developed inter-
metacarpal tubercle that did not reach the level of metacarpal III.
The pelvis had a small pneumatic fossa on the anterior portion of the
renal depression, and the pectineal process was small. The hypotarsus
had only one closed calcaneal canal, and the inner calcaneal ridge was
without a distal extension or spur core.
By Oligocene time the two basic types of pelvis had probably be-
come established, since the single known Oligocene fossil already re-
sembled specialized modern forms. By the Miocene fossa II of the
humerus was starting to develop, as shown by Miortyx teres. A vestige
of the inner shelf of the pneumatic fossa of the humerus was still pres-
ent in the middle Pliocene Lophortyx shotwelli.
The Odontophoridae are presently at the following stage of evolu-
tion: They are generally the smallest of the Galliformes. The rostrum
is relatively short and deep, with this condition reaching its culmina-
tion in Rhynchortyx. The sternum is deeply notched, with long, nar-
row, parallel anterior lateral processes; the pneumaticity of the an-
terior sternal plate and the dorsal manubrial foramen has been lost.
The coracoid has an overhanging ventral portion of the brachial tube-
rosity and a sharply raised dorsal intermuscular line; with one excep-
tion the termination of the ventral intermuscular line of the coracoid
has shifted to the tip of the sterno-coracoidal process, which now ends
in a terminal knob. The scapular blade is long and has a deep dorsal
groove. The humerus has a large pneumatic fossa, whose inner shelf
is completely lost, except in one species of Odontophorus; fossa II is
well developed in all genera but Dendrortyx and Odontophorus. The
carpometacarpus has a produced pisiform process, and the intermeta-
carpal tubercle extends to metacarpal III in all but a few individuals
of Cyrtonyx. The pelvis has lost the pneumatic fossa in the renal de-
pression, and in the Odontophorus group the pectineal process is
further reduced or even lost. Finally, the hypotarsus has developed
a second closed calcaneal canal in most genera.


Vol. 6






1961 HOLMAN: OSTEOLOGY OF QUAILS 209

In the best known genus, Colinus, a progressive trend is evident
from late Pliocene time to the present toward reduction in size and
gradual change in qualitative characters. Colinus hibbardi from the
upper Pliocene of Kansas is the most distinct species of Colinus on
the basis of its large size and qualitative osteological characters.
Colinus suilium from the pre-Wicomico Pleistocene of Florida is some-
what smaller than C. hibbardi and has fewer qualitative osteological
characters. Fossils of latest Pleistocene age are identical with living
C. virginianus, except for their slightly larger size. The change from
C. hibbardi to C. suilium apparently took place during the early Ple-
istocene. The transition from C. suilium to C. virginianus occurred in
Wicomico time and was probably initiated by the change from a
glacial to an interglacial climate.







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