A new species of Palaeortyx (Aves: Galliformes: Phasianidae) from the Neogene of Gargano, Italy

Abstract

A new species, Palaeortyx volans n. sp., of Phasianidae (Aves: Galliformes) is described from the Neogene vertebrate assemblage of the Gargano (Italy). The Gargano fossil vertebrate association has been well known since 1971, and it is considered to be an island fauna because of the high degree of endemism shown by different taxa. The original descrip-tion of the fossil avifauna of Gargano referred all the phasianid specimens to Palaeortyx grivensis, a fossil species described form the Miocene (MN7+8) of La Grive-Saint-Alban (France). The present study reveals the differences between Palaeortyx volans n. sp. and the other species of Palaeortyx described until now. The new species described herein represents the small-est species of the genus known, except for P. joleaudi from La Grive-Saint-Alban. The systematic position of P. grivensis from La Grive-Saint-Alban is supported. Morphometric analysis of the fossil remains of P. volans n. sp. suggests good flying capabilities and an ecology similar to that of the recent Coturnix coturnix. These features allowed P. volans n. sp. to colonize isolated islands such as Gargano, which are normally not inhabited by short-distance flyers like the phasianids. Une nouvelle espèce de Palaeortyx (Aves, Galliformes, Phasianidae) du Néogène du Gargano -Les gisements de Vertébrés néogènes de la presqu'île du Gargano (Italie) ont livré une nouvelle espèce de Phasianidae (Aves, Galliformes), décrite ici sous le nom de Palaeortyx volans n. sp.. Les faunes de ces gisements, signalées pour la première fois en 1971, sont des faunes insulaires dont les taxons présentent un fort degré d'endémisme. L'avifaune a été étudiée par Peter Ballmann en 1973 et 1976. Dans ces deux publications tous les restes de Phasianidae ont été attribués à Palaeortyx grivensis, une espèce éteinte décrite dans le Miocène (MN 7 + 8) de La Grive-Saint-Alban (France). Notre étude montre que la forme du Gargano est différente de toutes les autres espèces du genre Palaeortyx. La nouvelle espèce est la plus petite connue jusqu'à présent dans ce genre, à l'exception de P. joleaudi de La Grive-Saint-Alban. Notre étude confirme également la validité de l'espèce P. grivensis. L'analyse morphométrique des différents éléments de P. volans montre que cette espèce avait une bonne capacité de vol et suggère une autoécologie semblable à celle de l'actuelle Coturnix coturnix. Cette capacité lui a permis de coloniser des îles assez lointaines, comme celles de l'archipel du Gargano, qui normalement ne sont pas atteintes par les autres Phasianidae, qui volent seulement sur de faibles distances.

Full text

ORYCTOS vol. 7, 2008 95
A new species of Palaeortyx (Aves: Galliformes: Phasianidae) from
the Neogene of Gargano, Italy
Ursula B. Göhlich
1
& Marco Pavia
2
1
Naturhistorisches Museum Wien, geologisch-paläontologische Abteilung -- Burgring 7, A-1010 Wien. e-mail: ursula.
goehlich@nhm-wien.ac.at
Address when conducting the study: Humboldt-Fellow at the Université Claude Bernard - Lyon 1, UMR P.E.P.S., Bâtiment
Géode, 2 rue Dubois, F-69622 Villeurbanne Cedex, France.
2
Università di Torino - Museo di Geologia e Paleontologia, Dipartimento di Scienze della Terra, Via Valperga Caluso 35,
10125 Torino, Italy. e-mail: marco.pavia@unito.it
Corresponding author: Marco Pavia
ABSTRACT - A new species, Palaeortyx volans n. sp., of Phasianidae (Aves: Galliformes) is described from the Neogene
vertebrate assemblage of the Gargano (Italy). The Gargano fossil vertebrate association has been well known since 1971, and
it is considered to be an island fauna because of the high degree of endemism shown by different taxa. The original descrip-
tion of the fossil avifauna of Gargano referred all the phasianid specimens to Palaeortyx grivensis, a fossil species described
form the Miocene (MN7+8) of La Grive-Saint-Alban (France). The present study reveals the differences between Palaeortyx
volans n. sp. and the other species of Palaeortyx described until now. The new species described herein represents the small-
est species of the genus known, except for P. joleaudi from La Grive-Saint-Alban. The systematic position of P. grivensis
from La Grive-Saint-Alban is supported. Morphometric analysis of the fossil remains of P. volans n. sp. suggests good flying
capabilities and an ecology similar to that of the recent Coturnix coturnix. These features allowed P. volans n. sp. to colonize
isolated islands such as Gargano, which are normally not inhabited by short-distance flyers like the phasianids.
Keywords: Palaeortyx, Phasianidae, Galliformes, Gargano, Italy, Neogene,
Une nouvelle espèce de Palaeortyx (Aves, Galliformes, Phasianidae) du Néogène du Gargano - Les
gisements de Vertébrés néogènes de la presqu’île du Gargano (Italie) ont livré une nouvelle espèce de Phasianidae (Aves,
Galliformes), décrite ici sous le nom de Palaeortyx volans n. sp.. Les faunes de ces gisements, signalées pour la première fois
en 1971, sont des faunes insulaires dont les taxons présentent un fort degré d’endémisme. L’avifaune a été étudiée par Peter
Ballmann en 1973 et 1976. Dans ces deux publications tous les restes de Phasianidae ont été attribués à Palaeortyx grivensis,
une espèce éteinte décrite dans le Miocène (MN 7 + 8) de La Grive-Saint-Alban (France). Notre étude montre que la forme du
Gargano est différente de toutes les autres espèces du genre Palaeortyx. La nouvelle espèce est la plus petite connue jusqu’à
présent dans ce genre, à l’exception de P. joleaudi de La Grive-Saint-Alban. Notre étude confirme également la validité de
l’espèce P. grivensis. L’analyse morphométrique des différents éléments de P. volans montre que cette espèce avait une bonne
capacité de vol et suggère une autoécologie semblable à celle de l’actuelle Coturnix coturnix. Cette capacité lui a permis
de coloniser des îles assez lointaines, comme celles de l’archipel du Gargano, qui normalement ne sont pas atteintes par les
autres Phasianidae, qui volent seulement sur de faibles distances.
Mots clés: Palaeortyx, Phasianidae, Galliformes, Gargano, Italie, Néogène
INTRODUCTION
The fossil vertebrates of the Mio-Pliocene fissure
fillings of Gargano, Italy, are known since the 1970s (Freu-
denthal, 1971). The avifauna was first described by Ballmann
(1973, 1976); in these studies, all the galliform remains were
referred to Palaeortyx grivensis Lydekker, 1893. Several
studies on the mammal fauna [see Abbazzi et al. (1996) for
list of references] have been published that reveal that the
mammals represent an island fauna, based on the high de-
gree of endemism shown by some taxa. This fact can also
be observed in the avifauna (Ballmann, 1973, 1976; Pavia

ORYCTOS vol. 7, 2008 96
& Göhlich, 2004). The whole vertebrate association is very
rich, and it comprises several mammalian, avian, reptilian,
and amphibian taxa. It is known as the “Microtia fauna” after
the endemic murid rodent Microtia (Freudenthal, 1971). It
is assumed that the fissure fillings comprise a notable time
span. Whereas Freudenthal (1971, 1976) proposed a Middle
Miocene age for the deposits, after more recent analyses, Ab-
bazzi et al. (1996) came to the conclusion that the fossils rep-
resent a Late Miocene to Early Pliocene fauna. The “Microtia
fauna” is assumed to be the remnant of an endemic fauna that
inhabited a much larger bioprovince during the Miocene, an
interpretation that is also supported by the discoveries at the
Tortonian locality of Scontrone, Central Italy, where similar
taxa have been found (Mazza & Rustioni, 1996).
The fossiliferous localities of Gargano are
located in the area of Apricena (Foggia, Apulia, southern It-
aly) on the north-western slope of the Gargano promontory.
After the first report on the occurrence of fossil vertebrates
in this area (Freudenthal, 1971), systematic field research has
been carried out by the Nationaal Natuurhistorisch Museum
of Leiden (The Netherland) and the University of Firenze
(Italy). These field efforts established extensive and varied
collections of Neogene and Pleistocene vertebrate remains
(Freudenthal, 1976; De Giuli & Torre, 1984; Abbazzi et al.,
1996). The Pleistocene fossil assemblage consists of several
mammalian, avian, reptilian, and amphibian taxa, which
were described in several papers (Abbazzi et al., 1996, Delf-
ino & Bailon, 2000; Rook et al., 2004; Bedetti, 2003).
The Neogene fossil localities consist of a karst net-
work filled by residual clays (“terre rosse”) that are locally
very rich in continental vertebrate remains. These deposits
represent the “Microtia fauna”. The residual clays accumu-
lated over a wide area during a long period of emergence of
the Gargano region, probably from the Late Oligocene to the
Early Pliocene. Locally, these clays filled fissures opened in
the Mesozoic limestone following long-term sub-aerial ex-
posure of the carbonate platform (Abbazzi et al., 1996). The
fissure fillings, exposed in the walls of various quarries in the
area, are indicated by the name of the quarry or its owner,
followed by a number. As in other karst networks, one fis-
sure filling might be different in age from adjacent fissures,
thus the different fissures are treated as separate localities, as
listed in Ballmann (1973, 1976) and in the other systematic
papers on the Neogene Gargano fossil vertebrates (see Ab-
bazzi et al., 1996, for complete references).
MATERIAL AND METHODS
The material from Gargano described by Ballmann
(1973, 1976) is housed in the Naturalis – Nationaal Natu-
urhistorisch Museum, Leiden (The Netherlands), formerly
Rijksmuseum van Geologie en Mineralogie (RGM). This
paper concerns new material also housed at RGM, and also
several specimens in the collections of the Museo di Geolo-
gia e Paleontologia, University of Torino, Italy (PU), Dipar-
timento di Scienze della Terra, University of Firenze (IGF),
Figure 1 – Location map of the Apricena quarries area (left) and view of the Dell’Erba quarry (right top) and details on the extensive fos-
siliferous karst network developed in the Mesozoic limestone (right bottom).

ORYCTOS vol. 7, 2008 97
and the Naturmuseum Augsburg, Germany (NMA).
For this study, the specimens of Palaeortyx from
Gargano were compared with the type and additional materi-
al of Palaeortyx brevipes, P. gallica, P. grivensis, P. joleaudi,
and Palaeocryptonyx depereti from the following localities:
Saint-Gérand-le-Puy (Early Miocene, MN2, France), La
Grive-Saint-Alban (Middle Miocene, MN7+8, France), and
Quercy (Oligocene, MP25-28, France). This material is in
the collections of the Muséum National d’Histoire Naturelle,
Paris (MNHN), the Muséum d’Histoire Naturelle, Lyon
(ML), the Faculté des Sciences de Lyon (FSL) [= Université
Claude Bernard, Lyon 1 (UCBL)], and the Natural History
Museum, London (NHM). We also used for comparison ma-
terial of extant Coturnix coturnix and Coturnix coromandel-
ica from the collections of the Staatssammlung für Anthro-
pologie und Paläoanatomie München (SAPM) and UCBL.
Osteological terminology used in this publica-
tion principally follows Baumel et al. (1993) and occasion-
ally Ballmann (1969a, b); measurements were taken after
von den Driesch (1976). Anatomical abbreviations—cmc:
carpometacarpus; mc: metacarpale; tt: tibiotarsus; tmt: tar-
sometatarsus; dist.: distal; prox.: proximal.
Systematic Palaeontology
Class Aves Linnaeus, 1758
Order Galliformes Temminck, 1820
Family Phasianidae Vigors, 1825
Sub-family Phasianinae Vigors, 1825
Genus Palaeortyx Milne-Edwards, 1869
Synonymy—see Göhlich and Mourer-Chauviré
(2005).
The systematics of the species and the validity of
the genus Palaeortyx follow Göhlich & Mourer-Chauviré
(2005) and differ from those proposed by Mlíkovský (2000,
2002), which included a synonymy of Palaeortyx and Cotur-
nix (Mlíkovský, 2002: 152f).
Type species—Palaeortyx gallica Milne-Edwards,
1869 (designated by Milne-Edwards 1867-71: 230).
Diagnosis—A detailed generic diagnosis was given
by Ballmann (1969b: 30f), which he emended in 1973 (p.
24f). Cheneval (2000: 344) translated the original diagnosis
into French. Göhlich & Mourer-Chauviré (2005) presented
an English translation of Ballmann’s (1969b, 1973) original
and emended diagnoses, and further emended the diagnosis
of Palaeortyx.
Remarks—The phasianid represented in Gargano is
a very small sized species of Palaeortyx. Other small sized
Neogene phasianids, originally described as Palaeortyx are
P. brevipes Milne-Edwards, 1869, P. grivensis Lydekker,
1893, P. depereti Ennouchi, 1930, and P. joleaudi Ennouchi,
1930. Palaeortyx brevipes is recorded from its type locality
of Saint-Gérand-le-Puy (Lower Miocene, MN2) and several
other Late Oligocene and Early and Middle Miocene lo-
calities in Europe (see Göhlich & Mourer-Chauviré, 2005);
both P. grivensis and P. depereti have been described from
the Middle Miocene of La Grive-Saint-Alban. Ballmann
(1973, 1976) considered P. depereti as a junior synonym of
P. grivensis. However, the lectotype humerus of P. depereti,
selected recently by Mlíkovský (2002: 154), shows a shal-
low dorsal fossa pneumotricipitalis and a pneumatic ventral
fossa pneumotricipitalis, with a distal bony crest bordering
the fossa. Therefore Göhlich & Mourer-Chauviré (2005) re-
ferred it to Palaeocryptonyx Depéret 1892 [which is not to
be confused with Palaeocryptonyx depereti Gaillard (1908)
from Quercy, which was moved to Quercymegapodius
by
Mourer-Chauviré 1992: 81]. Palaeortyx grivensis from La
Grive-Saint-Alban was described by Lydekker in 1893 on
the basis of one humerus (NHM A319, fig. 2, A). Later,
Ennouchi (1930: 72) referred three more humeri (ML LGr
52-54) from the same locality to P. grivensis. We agree with
Ballmann (1969a, 1973, 1976), that most of the material in
the collections at ML and the FSL from La Grive locality
labelled as P. depereti cannot be distinguished in size and
morphology from P. grivensis. However, because Mlíkovský
(2002) later chose a specimen of P. depereti as lectotype,
which Göhlich & Mourer-Chauviré (2005) found to belong
to the genus Palaeocryptonyx, the species P. depereti and P.
grivensis cannot be synonymized as per Ballmann (1969a).
Furthermore, P. grivensis was synonymized with Coturnix
gallica by Mlíkovský (2002: 154) [actually P. gallica fol-
lowing Göhlich & Mourer-Chauviré (2005)], a decision with
which we do not agree because P. grivensis is clearly smaller
than P. gallica, as comparison with the type material dem-
onstrates. However, P. grivensis from La Grive-Saint-Alban
was later synonymized with P. brevipes from Saint-Gérand-
le-Puy by Göhlich & Mourer-Chauviré (2005). P. joleaudi,
known only by one humerus, is distinctly smaller than the
species from Gargano and all other named species and can
therefore be excluded from further comparisons.
Palaeortyx volans n. sp.
fig. 2, B-O
Synonymy:
Palaeortyx grivensis Lydekker – Ballmann 1973, p.
25, pl. 4, fig. 13-14
Palaeortyx grivensis Lydekker – Ballmann 1976, p.
8, pl. 3, fig. 1-9
Coturnix coturnix (Linnaeus) – Mlíkovský 2002,
p.155
Holotype – A complete left humerus RGM 178899,
from Chirò 27 (Ballmann, 1976) (fig. 2, B). The humerus
is chosen as the holotype because of the various species of
Palaeortyx are defined by their humeri (Göhlich & Mourer-
Chauviré, 2005).

ORYCTOS vol. 7, 2008 98

ORYCTOS vol. 7, 2008 99
Type locality – Chirò 27, Chirò quarry, Apricena,
(Foggia) Gargano area, Southern Italy (fig. 1). The Chirò
27 fissure excavaded by Freudenthal in the Chirò quarry
no longer exists because of the quarry activities; the Chirò
quarry is still active at the present time.
Paratypes – Right scapula RGM 425529, proxi-
mal half; left coracoid RGM 425513, almost complete; left
ulna RGM 425493, complete; left radaius RGM 425969,
complete; right carpometacarpus RGM 425504, lacking os
mc minus; left femur RGM 425490, complete; left tibiotar-
sus RGM 425501, distal half; right tarsometatarsus RGM
425514, lacking trochlea IV, all from the Chirò 27 fissure
filling.
Stratigraphic and geographic distribution – Palae-
ortyx volans n. sp. is reported from several fissure fillings of
the Gargano area, from the older (Biancone) to the younger
(San Giovannino) in the most recent interpretation of the
“Microtia fauna” evolution (Freudenthal, 1976).
Referred specimens – Material (see Annex).
Measurements – See table 1, 2 and 3.
Differential diagnosis – small sized Palaeortyx,
smaller than P. brevipes, but larger than P. joleaudi. Palae-
ortyx volans n. sp. differs from all other species of Palae-
ortyx by its wing and leg proportions, with a proportionally
longer carpometacarpus and a proportionally shorter tibio-
tarsus. Palaeortyx volans n. sp. differs from Coturnix co-
turnix and Palaeocryptonyx depereti (Ennouchi, 1930) by a
humerus with a deeper dorsal fossa pneumotricipitalis and
non-pneumatic ventral fossa pneumotricipitalis. Palaeortyx
volans n. sp. differs from Coturnix by a straighter humerus, a
carpometacarpus with craniocaudally longer ventral trochlea
carpalis and distally longer facies articularis digitalis minus,
a tibiotarsus with more strongly developed crista cnemialis
lateralis and cranialis, a scapula with a broader and more
angled acromion, a femur with a less developed crista tro-
chanteris, and a stouter tarsometatarsus.
Description and comparison – Palaeortyx volans
n. sp. from Gargano was compared with five specimens of
extant Coturnix coturnix (SAPM No. 3, female; SAPM No.
9, male, UCBL 130.1, 130.2, 130.4) and two specimens of
Coturnix coromandelica (UBCL
1980.1, 1980.2). This anal-
ysis was to detect any possible differences between these
forms and to verify the proposed synonymy of the Gargano
specimens with Coturnix coturnix as proposed by Mlíkovský
(2002: p. 155).
The scapula of Palaeortyx volans n. sp. differs from
that of Coturnix by having an acromion slightly broader
which in proximal view forms an angle of distinctly more
than 90° (about 110°) with the axis of the proximal end,
whereas it is a slightly thinner and forms an angle of about
90° in Coturnix. In the proximal part of the humerus, the
dorsal fossa pneumotricipitalis of Coturnix is more shallow
than in P. volans n. sp.; and the ventral fossa pneumotricipi-
talis is pneumatic in Coturnix, but not in P. volans n. sp.. The
humerus shaft of Coturnix is straighter and not curved as in
P. volans n. sp. (in caudal/cranial view). The proximal out-
line of the proximal end in P. volans n. sp. is characterized by
an indentation between the caput humeri and the tuberculum
ventrale (in caudal/cranial view), as in Perdix whereas this
indentation is very weak or lacking in Coturnix.
The carpometacarpus of P. volans n. sp. differs
from that of Coturnix by having the ventral side of the tro-
chlea carpalis more lengthened caudally and the distal facies
articularis digitalis minus more lengthened distally, whereas
in Coturnix the facies articulares digitales major and minor
have about the same length and are about the same level.
The crista trochanteris of the femur of Coturnix ex-
tends farther distally than in P. volans n. sp. Therefore, and
in contrast to Coturnix, the proximal end of the femur of P.
volans shows in lateral view a concavity of the cranial edge
distally to the crista trochanteris. In addition, the crista tro-
chanteris is proximally more protruding in P. volans, as can
be seen best in cranial view. Palaeortyx volans n. sp. shows
two foveae on the caput femoris, whereas there is only one
in Coturnix.
The proximal tibiotarsus of Palaeortyx volans n. sp.
differs from that of Coturnix by a stronger developed cristae
cnemialis lateralis and cranialis.
The tarsometatarsus of Palaeortyx volans n. sp. is
stouter than that of Coturnix.
As shown in fig. 3 the relationships of the limb bone
length in Palaeortyx volans n. sp. are different from that of
Coturnix coturnix. The humerus and femur of Palaeortyx vo-
lans n. sp. are shorter relative to that in C. coturnix, whereas
Figure 2 (opposite) – Long bones of Palaeortyx volans n. sp. from various locality of the Gargano area compared with the holotype of
P. grivensis from La Grive-Saint-Alban. A: Palaeortyx grivensis, holotype right humerus NHM A 319, caudal view, from La Grive-Saint-
Alban; B: Palaeortyx volans n. sp., holotype left humerus RGM 178899, caudal view, from Chirò quarry (Chirò 27); C: Palaeortyx volans
n. sp., distal left humerus RGM 425541, cranial view, from Chirò quarry (Chirò 14a); D: Palaeortyx volans n. sp., proximal left humerus
RGM 425519, caudal view, from Chirò quarry (Chirò 27); E Palaeortyx volans n. sp., paratype incomplete right scapula RGM 425529,
lateral view, from Chirò quarry (Chirò 27); F: Palaeortyx volans n. sp., paratype left ulna RGM 425493, ventral view, from Chirò quarry
(Chirò 27); G: Palaeortyx volans n. sp., left carpometacarpus PU 102029, ventral view, from Pirro quarry; H: same bone, dorsal view; I:
Palaeortyx volans n. sp., left coracoid RGM 425967, dorsal view, from Chirò quarry (Chirò 27); J: Palaeortyx volans n. sp., paratype left
femur RGM 425490, cranial view, from Chirò quarry (Chirò 27); K: same bone, caudal view; L: Palaeortyx volans n. sp., proximal right
tibiotarsus PU 102047, caudal view, from Dell’Erba quarry; M: Palaeortyx volans n. sp., paratype distal left tibiotarsus RGM 425501,
cranial view, from Chirò quarry (Chirò 27); N: Palaeortyx volans n. sp., left tarsometatarsus RGM 425553, dorsal view, from Chirò quarry
(Chirò 30c); O: same bone, plantar view. The scale bars represent 0,5 cm.

ORYCTOS vol. 7, 2008 100
the ulna is longer, even if the structure of Palaeortyx volans
n. sp. is more similar to that of Coturnix coturnix than to that
of other known species of Palaeortyx. The morphological
and metrical characteristics of Palaeortyx volans n. sp. indi-
cate that this species clearly belongs to the genus Palaeortyx,
as also shown by the shortness of the tarsometatarsus, which
is typical of the various species of this genus (fig. 3); it does
not belong to the modern genus Coturnix, particularly to C.
coturnix, as stated by Mlíkovský (2002).
DISCUSSION
In his papers on the Gargano fossil bird assemblag-
es, Ballmann (1973, 1976) referred all the galliform mate-
rial to Palaeortyx grivensis. This species was described by
Lydekker (1893) on the basis of a single humerus (NHM A
319) from La Grive-Saint-Alban, France. Later, Ennouchi
(1930) referred three more humeri to P. grivensis (ML LGr
52-54), and he described a second small sized species of
Palaeortyx from La Grive, P. depereti (Ennouchi 1930).
Following the opinion of Ballmann (1969a, 1973, 1976)
the latter is a junior synonym with P. grivensis. Recently,
Mlíkovský (2002) designated a lectotype for P. depereti, se-
lecting a complete humerus (ML LGr 50), that shows the
morphological characteristics of Palaeocryptonyx (Göhlich
& Mourer-Chauviré, 2005). For this reason, “depereti” must
be treated as a valid taxon in the genus Palaeocryptonyx and
therefore can no longer be considered as synonym of Palae-
ortyx grivensis
After evaluating the material, we agree with Ball-
mann that all of the Palaeortyx material from La Grive-
Saint-Alban belongs to one species. Recently Göhlich and
Mourer-Chauviré (2005) suggested that this species might
be a synonym of Palaeortyx brevipes. However, this study
supports the validity of P. grivensis in La Grive and dis-
tinguishes it from P. brevipes from Saint-Gérand-le-Puy,
Figure 3 – Ratio diagram of the log differences of the mean dimensions of the long bones of Palaeortyx volans n. sp. compared with
those of Coturnix coturnix (selected as standard), C. gomerae, C. delegorguei, C. coromandelica, Palaeortyx grivensis, P. phasianoides, P.
prisca, P. gallica, P. brevipes, Perdix perdix and Alectoris graeca (data from Kraft, 1974; Jaume et al., 1993; Göhlich & Mourer-Chauviré,
2005)

ORYCTOS vol. 7, 2008 101
as shown in fig. 4. To determine the validity of Palaeortyx
volans n. sp. we compared the holotype humerus (RGM
178899) with the holotype humerus of P. grivensis and the
referred humeri from La Grive-Saint-Alban. The humeri of
P. volans are morphologically and metrically very similar to
those of P. grivensis; however, the humerus of P. volans n.
sp. is somewhat smaller (fig. 4) and more slender; its proxi-
mal width and, especially, its distal width differ significantly
from those of P. grivensis (figs. 5, 6).
CONCLUSIONS
Previous work on the fossil bird material from Gar-
gano by Ballmann (1973, 1976) indicated Palaeortyx griven-
sis to be the only species of galliforms in the whole assem-
blage. However, the analyses presented here reveals that this
material belongs to the new species Palaeortyx volans n. sp..
This species is, up to now, only found in the Neogene Gar-
gano vertebrate assemblage, as shown in a recent revision
of the Miocene galliforms from France (Göhlich & Mour-
er-Chauviré, 2005) where no similar forms were detected.
Based on these data, P. volans n. sp. appears to be endemic
to that insular bioprovince.
However, limb proportions of different long bones
suggest that, compared to the other forms of the genus, Pal-
aeortyx volans n. sp. possessed good flying capabilities (fig.
3). In fact, the wing and leg and the relative dimensions of
the different bones are more similar to those of species of
the genus Coturnix, even if in Palaeortyx volans n. sp. the
shortness of the tarsometatarsus, typical of the genus Pal-
aeortyx, is evident. The genus Coturnix comprises a large
number of living species, widespread in different parts of the
world. Some of them show a good flying ability that allows
them to migrate, even over long distance, as the case of the
Common Quail, Coturnix coturnix, and to colonize islands,
even those far from the continent (Del Hoyo et al., 1994). In
general, galliforms are absent from islands, even from those
close to the mainland (Alcover et al., 1992; Mourer-Chau-
viré et al., 2001). If they are present, it indicates the end of
isolation of the island, as the case of the arrival of Alectoris
graeca in Sicily (Bonfiglio et al., 2002). The only confirmed
endemic species of galliform is Coturnix gomerae, from the
Pleistocene of Canary Islands. This species is clearly derived
from C. coturnix, and it shows wing and leg proportions that
clearly indicate a reduction of its flying ability (Jaume et al.,
1993).
The abundant fossil material of the new species
Palaeortyx volans n. sp. allows us to understand some of its
ecology. In fact, the presence of various remains of young
individuals, not yet able to fly and, at the same time, the
presence of medullary bone in some of the remains, clearly
indicate that this species was breeding in the Gargano area
(Ballmann, 1979; Matthiesen, 1990). The presence of a
breeding species on a island with no endemic characteristics
is normally interpreted as the proof of its migratory habit, for
this reason we suggest here that Palaeortyx volans n. sp. was
a migratory galliform, with an ecology very similar to the
living Coturnix coturnix, which is characterized by annual
migrations.
The good flying ability of Palaeortyx volans n. sp.
is the only way to explain its colonization of the Gargano
archipelago, probably quite far from the mainland, as dem-
onstrated by the high degree of endemism of the vertebrate
association. On the other hand the good flying ability sug-
Figure 4 – Total length of the humeri of small Palaeortyx: P. bre -
vipes from Saint-Gérand-le-Puy, P. grivensis and P. joleaudi from
La Grive-Saint-Alban, P. volans n. sp. from Gargano (data for P.
brevipes from Göhlich & Mourer-Chauviré, 2005).
Figure 5 – Value distribution of the proximal breadth of the hu-
merus in small Palaeortyx: P. brevipes from Saint-Gérand-le-Puy,
P. grivensis and P. joleaudi from La Grive-Saint-Alban, P. volans
n. sp. from Gargano (data for P. brevipes from Göhlich & Mourer-
Chauviré, 2005).

ORYCTOS vol. 7, 2008 102
gests that this species was not strictly endemic to the Gar-
gano area, in that case, in fact, the flying ability was expected
to be reduced, as observed in Coturnix gomerae. Palaeortyx
volans n. sp. can be expected to be found in some other fossil
assemblages of the Neogene, probably of the Mediterranean
area, as it was not recognised in France and Central Europe
(Göhlich & Mourer-Chauviré, 2005). In conclusion, this spe-
cies represents the most abundant bird taxon of the Gargano
Neogene vertebrate assemblages, where it was probably an
important food source for the numerous birds of prey, most-
ly strigiforms, present in the association (Ballmann, 1973,
1976; Pavia & Göhlich, 2004). On the other hand, Palae-
ortyx volans n. sp. indicates an open environment with low
vegetation and grassland for the Gargano archipelago during
the Neogene.
ACKNOWLEDGEMENTS
First of all we would like to thank in particular
M. Freudenthal (Leiden), F. Masini (Firenze), C. Petronio
(Roma) and M. Rummel (Augsburg) for putting the studied
material at our disposal. Special thank to L. van den Hoek
Ostende (Leiden) for the possibility to work on the material
published by Ballmann and housed in Leiden. We warmly
thank C. Mourer-Chauviré for continuous discussions and
suggestions and for translating the French abstract. Many
thanks are also given to K. Campbell (Los Angeles) for the
improvement of the English. We are grateful to S. Chapman
(London) and J. Cooper (Tring) for making the holotype of
P. grivensis available; photo of the holotype of P. grivensis
was made by P. Crabb of the Photographic Unit of the Natu-
ral History Museum, London. This research received sup-
port from the SYNTHESYS Project http://www.synthesys.
info/ which is financed by European Community Research
Infrastructure Action under the FP6 “Structuring the Euro-
pean Research Area” Programme, in particularly with NL-
TAF-2904 to M. Pavia; work supported by Italian MIUR
PRIN 2006 to G. Pavia. While conducting this study U. Göh-
lich was a Humboldt Fellow in the Feodor Lynen Program at
the Université Claude Bernard, Lyon 1, France. The authors
also thank C. Mourer-Chauviré and an anonymous referee
for helpful comments on the manuscript.
REFERENCES
Abbazzi, L., Benvenuti, M., Boschian, G., Dominici, S.,
Masini, F., Mezzabotta, C., Rook, L., Valleri, G. &
Torre, D. 1996. The Neogene and Pleistocene succession
and the mammal faunal assemblages of an area between
Apricena and Poggio Imperiale (Foggia). Memorie
della Società Geologica Italiana, 51: 383-402.
Alcover, J. A., Florit, F., Mourer-Chauviré, C. & Weesie, P. D.
M. 1992. The avifaunas of the isolated Mediterranean
islands during the Middle and Late Pleistocene; pp.
273-283. In Campbell, K. E. (ed.). Papers in avian
paleontology honoring Pierce Brodkorb. Science
Series, Natural History Museum of Los Angeles County,
36: 491 pp.
Ballmann, P. 1969a. Les oiseaux miocènes de La-Grive-
Saint-Alban (Isère). Geobios, 2: 157- 204.
Ballmann, P. 1969b. Die Vögel aus der altburdigalischen
Spaltenfüllung von Wintershof (West) bei Eichstätt in
Bayern. Zitteliana, 1: 5-60.
Ballmann, P. 1973. Fossile Vögel aus dem Neogen der
Halbinsel Gargano (Italien). Scripta Geologica, 17:
1-75.
Ballmann, P. 1976. Fossile Vögel aus dem Neogen der
Halbinsel Gargano (Italien) zweiter Teil. Scripta
Geologica, 38: 1-59.
Ballmann, P. 1979. Fossile Glareolidae aus dem Miozän des
Nördlinger Ries (Aves: Charadriiformes). Bonner
Zoologische Bieiträge, 30 (1-2): 53-101.
Baumel, J. J., King, A. S., Breazile, J. E., Evans, H. E.
and Van den Berge, J. C. 1993. Handbook of avian
anatomy: Nomina Anatomica Avium. Publication of
the Nuttall Ornithological Club, 23: 779 pp.
Bedetti, C. 2003. Le avifaune fossili del Plio-Pleistocene
italiano: sistematica, paleoecologia e elementi
di biocronologia. Unpublished PhD Dissertation
University “La Sapienza” of Roma, 246 pp.
Bonfiglio, L., Mangano, G., Marra, A. C., Masini, F., Pavia,
M. & Petruso, D. 2002. Pleistocene Calabrian and
Sicilian bioprovinces. Geobios, Mémoire special, 24:
29-39.
Cheneval, J. 2000. L’avifaune de Sansan; pp. 321-388. In
Ginsburg, L. (ed.). La faune miocène de Sansan et
son environnement. Mémoires du Muséum National
d’Histoire Natururelle de Paris, 183: 392 pp.
Figure 6 – Value distribution of the distal breadth of the humer-
us in small Palaeortyx: P. brevipes from Saint-Gérand-le-Puy, P.
grivensis and P. joleaudi from La Grive-Saint-Alban, P. volans n.
sp. from Gargano (data for P. brevipes from Göhlich & Mourer-
Chauviré, 2005).

ORYCTOS vol. 7, 2008 103
De Giuli, C. & Torre, D. 1984. A microfauna with
Allophaiomys pliocaenicus from Gargano (Southern
Italy). Palaeontographia Italica, 73: 116-128.
Delfino, M. & Bailon, S. 2000. Early Pleistocene herpetofauna
from Cava Dell’Erba and Cava Pirro (Apulia, southern
Italy). Herpetological Journal, 10: 95-110.
Del Hoyo, J., Elliott, A. & Sargatal, J. (eds) 1994. Handbook
of the Birds of the World. Vol. 2. New World Vultures to
Guineafowl. Lynx Edicion, Barcelona: 638 pp.
Depéret, C. 1892. Sur la faune d’oiseaux pliocènes
du Roussillon. Comptes Rendus des Séances
Hebdomadaires de l’Académie des Sciences (Paris,
114: 690-692.
Driesch, A. von den 1976. A guide to the measurements
of animal bones from archaeological sites. Peabody
Museum Bulletin, 1: 1-129.
Ennouchi, E. 1930. Contribution à l’Etude de la Faune du
Tortonien de La Grive St.-Alban (Isère). Les Presses
Modernes, Paris, 135 pp.
Freudenthal, M. 1971. Neogene vertebrates from the Gargano
Peninsula. Scripta Geologica, 3: 1-10.
Freudenthal, M. 1976. Rodent stratigraphy in some Miocene
fissure fillings in Gargano (prov. Foggia, Italy). Scripta
Geologica, 37: 1-23.
Gaillard, C. 1908. Les oiseaux des phosphorites du Quercy.
Annales de l’Université de Lyon, n .sér., 23: 178 pp.
Göhlich, U. B. & Mourer-Chauviré, C. 2005. Revision of
The Phasianids (Aves: Galliformes) from the Lower
Miocene of Saint-Gérand-le-Puy (Allier, France).
Palaeontology, 48(6): 1331-1350.
Jaume, D.; McMinn, M. & Alcover, J. A. 1993. Fossil
birds from the Bujero del Silo, La Gomena (Canary
Islands), with a description of a new species of Quail
(Galliformes: Phasianidae). Boletim do Museu
Municipal do Funchal, Sup. 2: 147-165. Kraft is cited
in figure caption
Kraft, E. 1972. Vergleichende morphologische
Untersuchungen an Einzelknochen nord-
und mitteleuropäischer kleiner Hühnervögel.
unpublished thesis, Institut für Paläoanatomie,
Domestikationsforschung und Geschichte der
Tiermedizin, University Munich: 194 p.
Linnaeus, C. v 1758. Systema naturae, British Museum of
Natural History (10th ed.), London, 824 pp.
Lydekker, R. 1893. On some Bird-bones from the Miocene
of Grive-St.-Alban, Department of Isère, France.
Proceedings of the Zoological Society of London, 35:
517-522.
Mazza, P. & Rustioni, M. 1996. The Turolian fossil
artiodactyls from Scontrone (Abruzzo, Central Italy)
and their paleoecological and paleogeographical
implications. Bollettino della Società Paleontologica
Italiana, 35: 93-106.
Matthiesen, D. G. 1990. Avian medullary bone in the
fossil record, an example from the early Pleistocene
of Olduvai Gorge, Tanzania. Journal of Vertebrate
Paleontology, 10 (3): 34A.
Milne-Edwards, A. 1867-1871. Recherches anatomiques et
paléontologiques pour servir à l’histoire des oiseaux
fossiles de la France, vol. 1 (1867-1868), vol.2
(1868-1871, see Zoological Record, vol. 5-8), 632 pp.,
Atlas 1+2 pls. 1-96 + pl. 97-200 [date of publication
of Palaeortyx gallica, and P. brevipes is 1869, see
Zoological Record, vol. 6. p. 93].
Mlíkovský, J. 2000. Early Miocene quails (Aves: Phasianidae)
from Saint-Gérand-le-Puy, France. Časopis Národního
Muzea Rada Přírodovědná, 169(1-4): 91-96.
Mlíkovský, J. 2002. Cenozoic birds of the world, part 1:
Europe. Ninox Press, Praha, 406 pp.
Mourer-Chauviré, C. 1992. The Galliformes (Aves) from
the Phosphorites du Quercy (France): Systematics
and biostratigraphy; pp. 67-95. In Campbell, K. E.
(ed.). Papers in avian paleontology honoring Pierce
Brodkorb. Science Series, Natural History Museum of
Los Angeles County, 36: 491 pp.
Mourer-Chauviré, C., Louchart, A., Pavia, M. & Segui, B.
2001. Les avifaunes du Pléistocène moyen et supérieur
des îles méditerranéennes. Bulletin de la Société des
sciences historiques et naturelles de la Corse, 696-697:
223-244.
Pavia, M. & Göhlich, U. B. 2004. Revision of the fossil bird
association of the Neogene of the Gargano (Apulia, SE
Italy). Abstracts of the 6
th
International Meeting of the
Society of Avian Palaeontology and Evolution, Quillan
(France): 50-51.
Rook, L., Martinez Navarro, B. & Howell, F.C. 2004.
Occurrence of Theropithecus sp. in the Late
Villafranchian of Southern Italy and implication for
Early Pleistocene ‘‘out of Africa’’ dispersals. Journal
of Human Evolution, 47: 267-277.
Temminck, C.J., 1820-1840. Manuel d’Ornithologie, ou
Tableau Systématique des Oiseaux qui se trouvent en
Europe;précédé d’une Analyse du Système général
d’Ornithologie, et suivi d’une table alphabétique des
Espèces.2nd ed., vol. I - II: i-cxv, 1-439 & 440-950-
Paris (Gabriel Dufour).
Vigors, N. 1825. Observations on the natural affinities that
connect the orders and families of birds. Transactions
of the Linnaean Society of London, 14, 395-517.
ANNEX – LIST OF EXAMINED MATERIAL OF
PALAEORTYX VOLANS N. SP.
Scapula—RGM 425952, left prox. half; RGM
425529, right prox half; RGM 425758, right prox. half;
RGM 425988, left prox. half; RGM 335994, left prox. half;
PU 102078, left prox. end.
Coracoid—PU 102045, left, lacking proc. latera-
lis; RGM 425513, left, almost complete; RGM 425729, left,
lacking proc. acrocoracoideum and proc. lateralis; IGF 112,
right, almost complete; RGM 425665, right, almost com-

ORYCTOS vol. 7, 2008 104
plete; RGM 425967, left, almost complete; RGM 425720,
right, lacking proc. lateralis; PU 102046, left, lacking
dist. end; RGM 425891, right, lacking the dist. end; RGM
425645, left, lacking dist. end; RGM 335982, left, lacking
proc. acrocoracoideum and proc. lateralis; RGM 425492,
left, lacking proc. acrocoracoideum; IGF 201, right, lacking
dist. end; RGM 425642, left, lacking dist. end; RGM 425667,
left, lacking dist. end; RGM 335981, left, lacking dist. end;
RGM 425560, right, lacking dist. end; RGM 425735, right,
lacking dist. end; RGM 425732, left, lacking dist. end; RGM
425524, left, lacking dist. end; RGM 335990, left, lacking
dist. end; RGM 425674, left, lacking dist. end; RGM 425997,
left, prox. half; RGM 335983, left, prox. half; RGM 425904,
right, prox. half; RGM 425677, right, prox. half; RGM
425630, right, prox. half; RGM 335992, right, prox. half;
RGM 425633, right, prox. half; RGM 425691, right, prox.
GL Lm Bp Dp Bd Did BF SC DC
Coracoid
IGF 112 20,8 20,0 6,2 2,6 - - 4,4 1,4 2,0
PU 102045 - 21,7 6,7 2,7 - - 5,0 1,6 2,1
RGM 424729 - 22,3* ----(4,5) 1,7 2,1
RGM 425513 Paratype - 20,2 6,7 2,3 - - - 1,5 2,1
RGM 425665 21,4 20,4 7,1 2,5 - - 4,7 1,7 2,2
RGM 425720 - 20,5 6,7 2,6 -
RGM 425967 22,7 22,1 7,2 2,6 - - 5,2 1,7 2,3
Humerus
PU 102022 32,5 - 8,1 - 5,7 - - 2,7 -
PU 102031 33,4 - 8,5 ----2,9
RGM 177582 31,1 - 7,7 - 5,3 - - 2,5 -
RGM 177615 29,5 - 7,0 - 5,2 - - 2,8 -
RGM 178676 32,4 - 7,9 - 5,4 - - 2,5 -
RGM 178757 30,7 - 7,4 - 5,4 - - 2,5 -
RGM 178897 31,4 - 7,6 - 5,5 - - 2,8 -
RGM 178899 Holotype 32,0 - 7,6 - 5,4 - - 2,7 -
RGM 178900 30,1 - 7,3 - 5,0 - - 2,7 -
RGM 178901 32,2 - 8,4 - 5,7 - - 2,7 -
RGM 178903 30,8 - 7,1 - 5,1 - - 2,6 -
RGM 178904 32,2 - 8,1 - 5,5 - - 2,6 -
RGM 178907 31,5 - 7,5 - 5,4 - - 2,4 -
RGM 178909 30,4 - 7,6 - 5,4 - - 2,5 -
RGM 178912 33,2 - 7,8 - 5,9 - - 2,7 -
RGM 178913 31,4 - 7,5 - 5,3 - - 2,6 -
RGM 178914 31,0 - - - 5,2 - - 2,6 -
RGM 178915 31,4 - 7,3 - - - 2,5 -
RGM 178916 31,7 - 7,9 - 5,3 - - 2,6 -
RGM 178918 31,2 - 7,6 - 5,2 - - 2,5 -
RGM 178924 31,6 - 7,9 - 5,7 - - 2,6 -
RGM 178925 31,1 - 7,6 - 5,5 - - 2,6 -
RGM 178930 30,5 - 7,3 - 4,9 - - 2,5 -
RGM 178964 33,5 - - - 5,5 - - 2,8 -
RGM 425850 32,6 - 7,7 - 5,7 - -
RGM 425851 31,7 - 7,8 ----2,7-
Ulna
IGF 026 31,6 - 3,5 4,9 2,7 3,1 - 1,5 3,7
PU 102027 26,3 - 3,5 5,0 3,3 3,2 - 1,5 3,8
PU 102069 26,5 - 3,3 4,5 3,0 2,9 - 1,4 3,5
RGM 425493 Paratype 29,8 - 3,4 4,9 3,2 3,2 - 1,5 3,7
RGM 425523 26,8 - 3,2 4,7 3,1 2,9 - 1,6 3,5
Radius
RGM 425969 Paratype 25,6 - 1,9 2,4 2,8 - - 1,1 -
Carpometacarpus
IGF 200 19,4 18,6 5,2 - - 3,7 - - -
PU 102029 19,7 18,7 5,2 - - 3,5 - - -
RGM 425504 Paratype - 16,2 4,6 - - - - -
RGM 425552 16,2 15,8 4,5 - - 3,1 - - -
RGM 425728 17,6 16,5 4,9 - - 3,2 - - -
RGM 425977 17,1 16,7 - - - 3,1 - - -
RGM 425985 17,3 16,9 4,8 - - 3,2 - - -
Table 1 –
Measurements of the complete bones of the forelimbs of Palaeortyx volans n. sp. Lm for the carpometacarpus is the length of
the os metacarpale major. * indicates estimated values.

ORYCTOS vol. 7, 2008 105
half; RGM 425983, right, prox. half; RGM 425958, left,
prox. half; RGM 425682, right, prox. half; RGM 335995,
right, prox. half; RGM 425745, right, prox. half; IGF 222,
right, prox. half; RGM 425559, left, prox. half; IGF 142,
left, prox. half; RGM 425746, left, prox. half; RGM 425989,
right, prox. half; RGM 425646, left, prox. half; IGF 36, left,
prox. half; RGM 425681, left, prox. half; RGM 425727, left,
prox. half; IGF 206, right, prox. end; IGF 65, left, prox. half;
RGM 335993, right, prox. half; RGM 425731, right, prox.
half; RGM 425738, right, prox. end; RGM 425717, right,
prox. half; PU 102079, right, prox. half; RGM 425683, left,
prox. half.
Humerus—PU 102022, right, complete; RGM
178897, right, complete; RGM 178900, right, complete;
RGM 178903, right, complete; RGM 178907, right, com-
plete; RGM 178916, right, complete; RGM 177582, left,
complete; RGM 177615, left, complete; RGM 178768, left,
complete; RGM 178757, left, complete; RGM 178899, left,
complete; RGM 178901, left, complete; RGM 178904, left,
complete; RGM 178909, left, complete; RGM 178912, left,
complete; RGM 178913, left, complete; RGM 178918, left,
complete; RGM 178924, left, complete; RGM 178925, left,
complete; RGM 178930, left, complete; RGM 425850, left,
complete; PU 102031, right, lacking epicondylus ventralis;
RGM 425851, left, almost complete; RGM 178929, right,
prox. part broken; RGM 178905, left, prox. part broken RGM
178908, left, prox. part broken; RGM 178914, left, prox. part
broken; RGM 178964, left, prox. part broken RGM 425854,
right, lacking dist. end; RGM 178921, right, prox. half;
RGM 178917, right, prox. half; RGM 178926, right, prox.
half; RGM 178902 right, prox. half; RGM 178967, right,
prox. half; RGM 178968, right, prox. half; RGM 177769,
right, prox. half; RGM 177749, right, prox. half; RGM
179018 right, prox. half; RGM 425845, left, prox. half; PU
102025, right, prox. half; IGF 204, right prox. half; RGM
425636, left, prox. half; RGM 425518, right, prox. end;
RGM 425890, right, prox. end; IGF 133, right, lacking prox.
end; RGM 425953, left, lacking prox. end; RGM 425815,
left, dist. half; RGM 425635, right, dist. half; RGM 425540,
right, dist. end; PU 102064, left, prox. half; RGM 425776,
left, prox. half; RGM 425889, right, prox. half, lacking ven-
tral tuberculum; PU 102062, right, prox. half; IGF 229, right,
prox. half; RGM 425503, right, prox. half; PU 102060, right,
prox. half; PU 102063, right, prox. half; PU 102061, right,
prox. half, lacking tuberculum ventrale; PU 102080, right,
prox. half, lacking tuberculum ventrale; RGM 425978, left,
prox. half, lacking tuberculum ventrale; IGF 70, left, prox.
half, lacking tuberculum ventrale; RGM 425516, left, prox.
end; RGM 425578, right, prox. end; RGM 425510, right,
prox. end; IGF 113, right, prox. end; RGM 425814, right,
prox. end; IGF 6, right, prox. end; RGM 178920, left, prox.
part; RGM 178911, left, prox. half; RGM 178910, left, prox.
part; RGM 178906, left, prox. half; RGM 178898, left, prox.
part; RGM 178896, left, prox. half; RGM 178895, left, lack-
ing dist. end; RGM 177699, left, prox. half; RGM 178942,
left, prox. part; RGM 177731, left, prox. half; RGM 178951,
left, prox. half; RGM 178922, left, prox. half; RGM 179010,
left, prox. part; RGM 425602, left, prox. end; RGM 425519,
left, prox. end; RGM 425520, left, prox. end; RGM 425855,
left, prox. end; RGM 335571, left, prox. half, lacking tuber-
culum ventrale; RGM 425530, left, prox. end; PU 102065,
right, lacking prox. end; IGF 139, left, lacking prox. end;
RGM 425783, left, lacking prox. end; IGF 189, right, lacking
prox. end; PU 102067, left, lacking prox. end; RGM 335984,
right, dist. half; RGM 425703, right, dist. half; IGF 12, right,
dist. half; RGM 335962, right, dist. half; IGF 100, right, dist.
half; RGM 425491, right, dist. half; PU 102068, right, dist.
half; RGM 425970, right, dist. half; RGM 425576, right,
dist. half; RGM 178927, right, dist. half; RGM 178947, right,
GL Lm Bp Dp Bd Did SC
Femur
PU 102026 35,0 32,4 6,3 3,7 5,8 5,0 2,4
PU 102081 33,8 32,1 5,9 3,8 5,7 4,8 2,5
RGM 425490 Paratype 34,1 32,1 6,3 4,1 5,7 (4,6) 2,4
RGM 425608 34,0 33,3 6,2 4,1 5,8 4,9 2,6
Tibiotarsus - -
IGF 061+057 41,2* 40,0* 4.8 6.3 - - 2.2
Tarsometatarsus - -
IGF 131 23,5 - - 4,0* - - 2,4
IGF 99 22,1* -----2,2
NMA 507/1801 23,9 - 5,0 4,9 - - 2,4
RGM 425514 Paratype 24,0 - 4,8 4,4 - - 2,4
RGM 425533 22,6 - 4,6* ---2,3
RGM 425553 22,2 - 4,6 4,3 5,0 - 2,2
RGM 425586 22,0 - 4,6 4,3 5,2 - 2,3
RGM 425778 24,5 - 5,1 4,8 5,4 - 2,5
RGM 425817 25,0 - 4,9 4,7 - - 2,4
RGM 425968 22,4 - 4,1* ---2,4
Table 2 –
Measurements of the complete bones of the hindlimbs of Palaeortyx volans n. sp. * indicates estimated values.

ORYCTOS vol. 7, 2008 106
Table 3 – Mean values of the greatest length (GL) and internal length (Lm, only for the coracoid) of the long bones of Coturnix coturnix,
C. gomerae, C. delegorguei, C. coromandelica, Palaeortyx volans n. sp., P. grivensis, P. phasianoides, P. prisca, P. gallica, P. brevipes,
Perdix perdix, Alectoris graeca and the values of the log differences with Coturnix coturnix, selected as standard (data from Kraft, 1974;
Jaume et al., 1993; Göhlich & Mourer-Chauviré, 2005).
C
ORACOID HUMERUS ULNA CARPOMETACARPUS FEMUR TIBIOTARSUS TARSOMETATARSUS
Taxon
x Lm n° log diff. x GL n° log diff. x GL n° log diff. x GL n° log diff. x GL n° log diff. x GL n° log diff. x GL n° log diff.
Coturnix
coturnix
23,7 7 0 32,8 7 0 28,2 8 0 17,9 7 0 35,8 7 0 42,4 7 0 26,2 7 0
Coturnix
gomerae
22,7 2 0,016 35,8 5 0,038 29,1 4 0,014 - - - 40,9 12 0,058 48,8 5 0,061 27,3 5 0,018
Coturnix
delegorguei
21,6 1 -0,006 33,7 1 0,012 29,7 1 0,023 - - - 35,3 1 -0,006 43,6 1 0,012 26,5 1 0,005
Coturnix
coromandelica
21,0 2 -0,018 32,9 5 0,001 27,6 3 -0,009 17,3 2 -0,015 32,6 3 -0,041 40,9 2 -0,016 24,7 3 -0,026
Palaeortyx
volans n. sp.
21,7 9 -0,004 31,6 26 -0,017 28,5 6 0,005 17,6 12 -0,006 34,0 12 -0,022 41,4 4 -0,010 22,8 18 -0,060
Palaeortyx
grivensis
24,4 1 0.013 34,0 5 0,014 - - - - - - 37,7 2 0,023 - - - - - -
Palaeortyx
phasianoides
37,5 4 0,234 53,3 6 0,211 (51) 1 0,257 29,2 1 0,213 53,2 1 0,172 70,3 2 0,219 - - -
Palaeortyx
prisca
33,5 4 0,185 49,3 3 0,177 47,7 2 0,228 - - - 47,9 7 0,126 68,3 1 0,207 37,1 1 0,151
Palaeortyx
gallica
30,0 3 0,137 41,1 9 0,098 38,7 5 0,138 - - - 42,1 7 0,070 59,8 3 0,149 33,0 3 0,100
Palaeortyx
brevipes
24,4 3 0,047 36,4 5 0,045 32,7 3 0,064 18,7 1 0,019 37,9 1 0,025 50,1 1 0,073 28,8 3 0,041
Perdix
perdix
34,9 62 0,202 48,8 48 0,172 44,5 47 0,198 26,9 49 0,176 55,6 61 0,191 70,3 46 0,222 41,2 48 0,197
Alectoris
graeca
38,0 26 0,239 52,6 21 0,205 48,8 20 0,238 30,4 20 0,230 61,5 23 0,235 81,9 20 0,286 45,8 19 0,246

ORYCTOS vol. 7, 2008 107
dist. half; RGM 178928, right, dist. half; RGM 179019,right,
dist. half; RGM 178704, right, dist. half; RGM 425532,
left, dist. half; IGF 121, left, dist. half; RGM 335969, left,
dist. half; IGF 134, left, dist. half; RGM 425640, left, dist.
half; RGM 335972, left, dist. half; RGM 425521, left, dist.
half; RGM 425580, left, dist. half; RGM 178898, left, dist.
half; RGM 178933, left, dist. half; RGM 177603, left, dist.
half; RGM 177602, left, dist. half; RGM 177729, left, dist.
part; RGM 177644, left, dist. part; RGM 177645, left, dist.
part; RGM 177621, left, dist. part; RGM 178919, left, dist.
half; RGM 425541, left. dist. end; RGM 425670, left. dist.
end; RGM 425696, left. dist. end; IGF 44, left. dist. end;
RGM 425893, left. dist. end; IGF 135, left. dist. end; RGM
425663, left. dist. end; RGM 425964, left. dist. end; RGM
425902, right, dist. end; RGM 425790, right, dist. end; IGF
120, left, lacking prox. end; IGF 29, right, prox. shaft; RGM
425708, left shaft; RGM 178923, left shaft; RGM 177654,
left shaft; RGM 425951, left shaft; IGF 211, left shaft; IGF
224, left shaft.
Ulna— IGF 26, right, complete; PU 102027, right,
complete; RGM 425493, left, complete; RGM 425523, left,
complete; PU 102069, left, complete; PU 102030, left, prox.
half; RGM 425796, right, prox. end; IGF 80, right, dist. half;
RGM 425971, left, prox. and dist. ends damaged; IGF 87,
left, lacking dist. end and acromion; RGM 425581, left,
prox. end; IGF 81, left, dist. half; RGM 335987, left, dist.
half; RGM 425789, right, dist. half; IGF 147, left, dist. half;
RGM 425484, right, dist. end; RGM 425747, right, dist. end;
RGM 335991, right, dist. end; PU 102070, left, proximal
half, lacking the acromion.
Radius—RGM 425969, left, complete.
Carpometacarpus—PU 102029, left, lacking os
mc minus; IGF 200, right, lacking os mc minus; RGM
425728, left, lacking os mc minus; RGM 425552, left, lack-
ing os mc minus; RGM 425504, right, lacking os mc minus;
RGM 335977, right, lacking os mc minus; RGM 425985,
right, lacking os mc minus; IGF 146, left, prox. end dam-
aged, lacking os mc minus; RGM 425517, left, lacking proc.
extensorius and os mc minus, dist. end damaged; RGM
425898, right, dist. end damaged, lacking os mc minus; RGM
425675, right, lacking os mc minus and the dist. end; RGM
425998, left, prox. half, lacking proc. extensorius, os mc mi-
nus; RGM 425896, left, prox. half, lacking os mc minus; IGF
187, left, proximal half, lacking the trochlea carpalis and os
mc minus; RGM 425736, right, prox. half, lacking os mc
minus; RGM 425858, right, lacking prox. end and os mc mi-
nus; RGM 425737, right, prox. half, prox. end damaged and
lacking os mc minus; RGM 425692, right, prox. half, prox.
end damaged and lacking os mc minus.
Femur—RGM 425608, left, complete; PU 102026,
right, complete; RGM 425490, left, complete; RGM 425607,
right, almost complete; RGM 425785, right prox. half; IGF
193, right, prox. half; RGM 335967, left, dist. 2/3; RGM
425792, right, dist. end; PU 102081, left, complete; PU
102071, right, prox. half, prox. end slightly damaged; IGF
136, right, prox. half; IGF 25, right, damaged prox. half;
RGM 425712, right, damaged prox. half; RGM 425981, left,
damaged prox. half; RGM 425566, left, damaged prox. half;
RGM 425575, left, damaged prox. half; RGM 425724, left,
damaged prox. half; RGM 425959, right, prox. half; RGM
425960, left, damaged prox. half; RGM 425525, left, prox.
half; RGM 425754, left, damaged prox. half; RGM 425721,
left, damaged prox. half; RGM 425714, left, damaged prox.
half; RGM 425973, right, damaged prox. half; PU 102082,
right, prox. half; RGM 425511, right, prox. half; RGM
425900, right, prox. end; PU 102073, right, dist. half; PU
102072, right, lacking the prox. end; RGM 425705, left, dist.
half; RGM 425515, left, dist. half; IGF 122, left, dist. half;
RGM 425730, left, dist. half; RGM 425568, right, dist. half;
IGF 186, right, dist. half; RGM 425573, left, dist. end; IGF
118, left, dist. end; IGF 140, left, dist. end; RGM 425962,
right, dist. end; IGF 35, right, dist. end; RGM 425871, right,
dist. end; RGM 425874, right, dist. end; RGM 425966, left,
shaft; RGM 425666, right, dist. end; RGM 425899 right,
dist. end.
Tibiotarsus—PU 102024, right, lacking prox. end;
IGF 61+57, right, complete; PU 102047, right, prox. end;
RGM 335978, left, dist. half; RGM 425509, right, prox. 2/3;
RGM 335976, left, prox. half; IGF 194 left, prox. end; RGM
425631, left, prox. end; RGM 335960, left, prox. end; RGM
425563, left, prox. end; RGM 425711, left, prox. end; RGM
335964, left, dist. half; PU102074, right, dist. half; IGF 108,
right, dist. half; IGF 145, left, dist. half, lacking condylus
medialis; IGF 91, left, dist. half; RGM 425690, left, dist.
half; RGM 425527, left, dist. end; RGM 425744, right, dist.
end; IGF 141, left, dist. end; RGM 425956, left, dist. end;
RGM 335978, right, dist. end; RGM 425924, right, dist. end;
RGM 425938, right, dist. end; RGM 425808, left, dist. end;
RGM 335968, right, dist. end; RGM 425980, left, dist. end;
RGM 425673, right, dist. end; RGM 425522, left, dist. end;
RGM 425671, right, dist. end; RGM 335974, right, dist. end;
RGM 425528, right, dist. end; RGM 425722, left, dist. end;
RGM 425488, right, dist. end; RGM 425995, left, dist. end;
RGM 425680, left, dist. end; RGM 425501, left, dist. end;
IGF 178, right, dist. end; RGM 425784, right, dist. end; IGF
167, right, dist. end; RGM 425644, left, dist. end; IGF 67,
right, dist. end; RGM 425498, right, dist. end; RGM 425852,
right shaft.
Tarsometatarsus—RGM 425778, right, complete;
RGM 425992, left, complete; RGM 425817, right, lacking
trochlea IV; RGM 425514, right, lacking trochlea II; RGM
425533, right, almost complete; RGM 425968, right, lacking
trochlea II, prox. end damaged; IGF 131, right, lacking tro-
chlea II, prox. end damaged; RGM 425553, left, complete;
RGM 425586, right, almost complete; PU 102028, left, dist.
half; IGF 99, left, lacking trochlea IV, prox. end damaged;

ORYCTOS vol. 7, 2008 108
PU 102076, right, lacking prox. end and trochleae II and
IV; PU 102075, left, lacking dist. end; RGM 425506, right,
shaft; IGF 97, right, lacking dist. end; RGM 425632, right,
lacking trochleae III and IV; RGM 425799, left, lacking dist.
end; RGM 335985, left, prox. half; RGM 425669, left, prox.
half; IGF 182, left, prox. half; RGM 425647, left, prox. half;
RGM 335975, right, prox. half, prox. end damaged; RGM
425993, right, prox. half; RGM 425972, right, dist. half,
lacking trochlea IV; RGM 425707, right, dist. half; RGM
425986, right, dist. half, lacking trochlea II; RGM 425939,
right, dist. half; RGM 425788, right, dist. half, lacking tro-
chlea IV; RGM 425963, right, dist. end; RGM 425679, right,
dist. end, lacking trochlea IV; RGM 425716, right, dist. end,
lacking trochlea IV; RGM 425957, right, dist. end, lacking
trochleae II and IV; RGM 425987, right, dist. end, lacking
trochlea II; RGM 425557, left, dist. half, lacking trochlea IV;
RGM 425734, left, dist. end; RGM 425994, left, dist. half,
lacking trochlea III; RGM 425892, left, dist. half, lacking
trochleae II and III; RGM 335986, left, dist. half, lacking
all trochleae; NMA 507/1801, left, almost complete, lacking
trochlea IV.
Sternum—RGM 425485, cranial half; RGM
425496, cranial half; RGM 425507, cranial portion; IGF
102, cranial fragment; IGF 198, cranial fragment.
Furcula—RGM 425787, fragment with apophysis
furculae.