A nearly complete skeleton of the fossil galliform bird Palaeortyxfromthe late Oligocene of Germany

INTRODUCTION

As traditionally recognized (e.g. Stresemann 1927–1934), the galliform taxon Phasianidae includes the Odontophorinae (New World quails), Numidinae (guineafowl), Meleagridinae (turkeys), Tetraoninae (grouse) and the, probably para- phyletic, “Phasianinae” (Old World quails, pheas- ants, partridges, etc.; Kimball et al. 1999, Arm- strong et al. 2001). Of these, only Tetraoninae and

“Phasianinae” occur in Europe today.

Although the European fossil record of galli- form birds dates back into the early Eocene (47 million years ago, Mayr 2000, 2006), all Eocene and early Oligocene galliform taxa are outside crown group Galliformes and belong to the Gallinuloi- didae, Quercymegapodiidae, and Paraortygidae (Mourer-Chauviré 1992, Mayr 2005). Phasianidae do not occur in Europe before the end of the early Oligocene, about 30 million years ago (Mourer- Chauviré 1992), and most Oligocene phasianids belong to Palaeortyx Milne-Edwards, 1869. This taxon also occurs in the early Miocene of France and Germany (Göhlich & Mourer-Chauviré 2005) and has so far been known from isolated limb and

pectoral girdle bones only. The only other phasianid from the Oligocene of Europe is Schaubortyx keltica (Eastman 1905), which is based on a rather poorly preserved disarticulated skele- ton from the early Oligocene (MP 25) of France (Eastman 1905, Schaub 1945) and distinctly differs from Palaeortyx in its intermembral proportions (Mourer-Chauviré 1992; contra Mlíkovský 2002 who synonymized Schaubortyx Brodkorb, 1964 and Palaeortyx Milne-Edwards, 1869).

Here we describe a nearly complete and artic- ulated skeleton of Palaeortyx from the late Oligo- cene (MP 28, Storch et al. 1996; i.e. 24.7 mya; Mertz et al. in press) maar lake deposit of Enspel near Bad Marienberg in Westerwald, Germany. It is the second avian specimen from Enspel (Mayr 2001), and one of a few articulated skeletons of Oligo- cene galliform birds.

MATERIAL AND METHODS

Osteological terminology follows Baumel &

Witmer (1993). Measurements are in millimeters and indicate the maximum length of the bone

A nearly complete skeleton of the fossil galliform bird Palaeortyx from the late Oligocene of Germany

Gerald M

, Markus P

& Michael W

1Forschungsinstitut Senckenberg, Sektion für Ornithologie, Senckenberganlage 25, D-60325 Frankfurt am Main, GERMANY, e-mail: Gerald.Mayr@senckenberg.de

2Landesamt für Denkmalpflege Rheinland-Pfalz, Referat Erdgeschichtliche Denkmalpflege, Große Langgasse 29, D-55116 Mainz, GERMANY, e-mail: erdgeschichte@landesdenkmalamt.rlp.de

Mayr G., Poschmann M., Wuttke M. 2006. A nearly complete skeleton of the fossil galliform bird Palaeortyx from the late Oligocene of Germany. Acta Ornithol. 41: 129–135.

Abstract. Phasianid galliform birds do not occur in Europe before the end of the early Oligocene, and their Paleogene fossil record mainly comprises isolated bones. Here we describe a nearly complete and articulated skeleton of a phasianid galliform bird from the late Oligocene of Enspel in Germany. The specimen is assigned to Palaeortyx cf.

gallica Milne-Edwards, 1869 and for the first time allows the recognition of cranial and pelvic details in a late Oligocene phasianid. Many gizzard stones are preserved in both the gizzard and the crop, and constitute the earliest fossil evidence for grit use in a phasianid galliform.

Key words: fossil birds, Galliformes, Palaeortyx, gizzard stones, Oligocene Received — July 2006, accepted — Oct. 2006

along its longitudinal axis. The specimen was pre- pared with the resin transfer method (Kühne 1961).

SYSTEMATIC PALEONTOLOGY Galliformes Temminck, 1820

Phasianidae sensu Stresemann (1927–34) Palaeortyx cf. gallica Milne-Edwards, 1869

Referred specimen. PW 2005/5023a-LS (Figs. 1–4) and PW 2005/5023b-LS (few bones only, not fig- ured), currently deposited in Landesamt für Denkmalpflege Rheinland-Pfalz, Mainz (the spec- imen will be transferred to the Landessammlung für Naturkunde RLP, Germany).

Locality and horizon. Enspel near Bad Marien- berg in Westerwald, Rheinland-Pfalz, Germany;

late Oligocene, MP 28 (Storch et al. 1996).

Dimensions (in brackets those of Palaeortyx gallica

Fig. 1. Palaeortyx cf. gallica Milne-Edwards, 1869, articulated skeleton from the late Oligocene of Enspel, Germany (PW 2005/5023a- LS) with interpretative drawing. The arrows indicate the areas where grit is accumulated. Abbreviations: fur — furcula, lcm — left carpometacarpus, lco — left coracoid, lfe — left femur, lhu — left humerus, ltb — left tibiotarsus, ltm — left tarsometatarsus, lul — left ulna, pel — pelvis, rcm — right carpometacarpus, rhu — right humerus, rtb — right tibiotarsus, rtm — right tarsometatarsus, rul — right ulna, sk — skull, ste — sternum. Specimen coated with ammonium chloride. Scale bar equals 20 mm.

as given by Göhlich & Mourer-Chauviré 2005).

Skull, 48.7. Beak, length from nasofrontal hinge to tip, ~19.1. Synsacrum, 42.1. Coracoid, ~33 [~31].

Humerus, 44.2 (left), 45.2 (right) [38.9–43.2]. Ulna,

~42–43 [38.4–40.5]. Carpometacarpus, 24.3 (left), 24.1 (right) [-]. Femur, ~42–43 (left) [39.1–44.2].

Tibiotarsus, 61.1 (left), 61.4 (right) [58.6–60.8]. Tar- sometatarsus, 34.4 (left), ~34.7 (right) [31.7–34.4].

Description and comparisons. The major wing and hindlimb elements of Palaeortyx were already described in detail (Milne-Edwards 1867–1871, Ballmann 1969, Mourer-Chauviré 1992), and thus the following description focuses on previously unknown skeletal elements.

The skull of Palaeortyx has so far been un- known and the Enspel galliform actually provides the first skull details of an Oligocene phasianid (Fig. 3), the skull of Schaubortyx being very poorly preserved. The beak measures about one third of the entire skull length and resembles that of the Common Partridge Perdix perdix in its proportions;

it has a rounded tip and is not as short and strong- ly decurved as the beak of the Odontophorinae (Holman 1964). The processus maxillaris of the os nasale, which borders the caudal end of the large narial opening (Fig. 3), is wide as in the Cracidae (guans, chachalacas, currasows) and Numidinae, wider than in most “Phasianinae” (except, for example, Afropavo and Pavo). The os praefrontale is well-developed and similar in shape to the corre- sponding bone of P. perdix; this bone is more reduced in, e.g., Coturnix, Rollulus, and Ammo- perdix, and articulates in a recess of the os frontale in Numidinae. As in other Phasianidae, the de- scending process of the os praefrontale is greatly reduced and does not reach the jugal bar.

Although the left sclerotic ring is completely pre- served, the exact number of ossicles cannot be counted but may have been 14 or 15 as in extant Galliformes. The processus postorbitalis is well developed as in other Phasianidae (Fig. 3); this process is very short in the Australasian Mega- podiidae (brush-turkeys, scrubfowl, megapodes) and the Gallinuloididae (Mayr & Weidig 2004).

The processus zygomaticus, however, appears to have been short, whereas it is very long and fuses with the processus postorbitalis via an ossified aponeurosis zygomatica in Cracidae and most Phasianidae; only in Numidinae it is vestigial. As in other Phasianidae except for the short-billed Odontophorinae, the processus orbitalis of the quadratum is elongated and slender (Fig. 3). The mandible bears a well-developed processus retro- articularis; a fenestra mandibulae is absent (in

Fig. 2. Palaeortyx cf. gallica Milne-Edwards, 1869, articulated skeleton from the late Oligocene of Enspel, Germany (PW 2005/5023a-LS); photo taken before the fossil was transferred onto the artificial matrix. A dark halo representing fossilized feathers is clearly visible. Scale bar equals 10 mm.

contrast to, e.g., Tetraoninae in which it is very large). The well-preserved hyoid apparatus (Fig.

3), which is also visible through the reverse of the transparent slab, matches that of extant phasian- ids.

The morphology of the cervical vertebrae cor- responds to that of extant Phasianidae. As in the latter, the third and fourth cervical vertebrae exhibit an osseous bridge from the processus transversus to the processus articularis caudalis (Fig. 3). Also as in extant Phasianidae, there are 20 praesacral vertebrae, and there is one free thoracic vertebra between the synsacrum and the notari- um (part of which can be seen in PW 2005/5023b- LS). The tail vertebrae are not preserved.

The facies articularis scapularis of the coracoid is shallow (PW 2005/5023b-LS) which is a de- rived characteristic of crown group Galliformes (Mourer-Chauviré 1992, Mayr 2000); the dorsal surface of the extremitas sternalis does not exhibit

Fig. 3. Palaeortyx cf. gallica Milne-Edwards, 1869, skull (PW 2005/5023a-LS) in dorsolateral (A) and ventrolateral (B) views with interpretative drawings. The latter photo was taken before the fossil was transferred into the artificial resin. Abbreviations: bhy

— basihyale, cbr — ceratobranchiale, fmg — foramen magnum, jug — os jugale, mnd — mandible, pgl — paraglossum, ppo — processus postorbitalis, prf — os praefrontale, qdr — quadratum, ret — processus retroarticularis, scl — sclerotic ossicles, uhy — urohyale. The cervical vertebrae are numbered. Specimen in (A) coated with ammonium chloride. Scale bars equal 5 mm.

pneumatic openings. The preservation of furcula and scapula prevents recognition of osteological details.

The sternum of Palaeortyx so far has not been described and is visible from its right side in PW 2005/5023a-LS. The carina sterni is well-devel- oped, with a pointed apex carinae and a convex cranial margin which bears a marked sulcus cari- nae. The incisions in the caudal margin are very deep as in extant Odontophorinae and “Phasia- ninae”, although it is difficult to discern the tra- beculae (the trabecula intermedia extends along a crack in the original slab, the caudal portion of the trabecula lateralis overlays the cranial end of the synsacrum). Details of the cranial portion of this bone, including the shape of the spina externa, are not visible.

The caudal surface of the proximal humerus of Palaeortyx is very characteristic owing to the pres- ence of a marked, double fossa pneumotricipitalis.

Unfortunately, this feature is not clearly visible in

the Enspel galliform, although a part of the caudal surface of the proximal humerus was exposed before embedding in the resin and can still be seen through the reverse of the transparent slab.

The carpometacarpus exhibits a processus inter- metacarpalis (Fig. 4) which is a derived feature of the Phasianidae and absent in Paleogene stem group Galliformes; this process is secondarily reduced in Numidinae (Stegmann 1978). As in many extant Phasianidae (e.g. Alectoris chukar), there is an ossified tendon along the craniodorsal surface of the carpometacarpus. The phalanx proximalis digiti majoris is not as abbreviated and craniocaudally wide as in some extant “Phasia- ninae” (e.g., Rollulus and Lophura). The phalanx digiti alulae bears a small ungual phalanx which is comparable in size to that of Alectoris.

The pelvis appears to have been of similar pro- portions to that of most extant Phasianidae and not as wide as in the Tetraoninae, although its original proportions are difficult to reconstruct

because of to the flattening of the specimen. Most notably, the tubercula praeacetabularia are very small as in extant Odontophorinae (Holman 1964), Tetraoninae, and a few taxa within “Phasia- ninae”, such as Ammoperdix and, according to Dyke et al. (2003), Arborophila and Galloperdix (Fig.

4); they are much more developed in other Phasianidae.

As in Palaeortyx, the femur is subequal to the humerus in length, whereas it is much longer in Schaubortyx keltica, extant “Phasianinae”, and most Odontophorinae. The femur is shorter than or subequal to the humerus in stem group Gal- liformes and extant Cracidae, Numidinae, Cyr- tonyx (Odontophorinae; Holman 1961), and Tetraoninae. Only the distal end of the tibiotarsus shows osteological details and resembles the distal tibiotarsus of Palaeortyx. The tarsometatarsus also agrees with that of Palaeortyx in its proportions and morphology. It lacks a spur which within extant Phasianidae is also wanting in the males of Odontophorinae, Numidinae, and Tetraoninae (contra Dyke et al. 2003), as well as Perdix (contra Dyke et al. 2003), Margaroperdix, Melanoperdix, Coturnix, Ammoperdix, Rollulus, Arborophila (contra Dyke et al. 2003), Xenoperdix, and Ptilopachus.

As in other Phasianidae, the hallux is pro- cumbent and more elevated than in Mega- podiidae and Cracidae; its ungual phalanx is very short. An ossified tendon attaches to the processus articularis tarsometatarsalis of the os metatarsale I and extends along the plantar surface of the tarsometatarsus.

Although remains of the primaries of the right wing are preserved, they do not permit a mean- ingful description. Ornamental feathers (e.g., on the head) cannot be discerned.

A large quantity of grit is preserved in a well- defined area of about 21 x 28.5 mm, between the pelvis and the sternum, and unquestionably represents gizzard stones (Figs. 1 and 4). A fewer amount of grit is also visible in the crop area, next to the proximal end of the right humerus (Fig. 1).

Grit size varies between 0.1 and 2.6 mm, on average the particles have a diameter of 1.5 mm;

all are rounded. Most of the gizzard stones are white and grayish quartz pebbles, about one fifth is reddish pyroclastic material whereas almost all grit particles in the crop area are reddish.

A brown organic substance with a fibrous texture suggestive of plant material is preserved in some places between the gizzard stones. Some unidentifiable remains of seeds are preserved in the crop area.

Fig. 4. Palaeortyx cf. gallica Milne-Edwards, 1869 (PW 2005/5023a-LS); distal part of the left wing (A) and pelvis with gizzard stones (B). Abbreviations: pim — processus inter- metacarpalis, tpa — tuberculum praeacetabulare. Specimen coated with ammonium chloride. Scale bars equal 5 mm.

DISCUSSION

The derived presence of a processus inter- metacarpalis on the carpometacarpus supports classification of the Enspel galliform into Phasia- nidae. It is assigned to Palaeortyx, the most abun- dant phasianid taxon in the late Oligocene of Europe, because of its intermembral proportions with the femur being as long as the humerus. The preservation does not permit detailed osteological comparisons, but in size and morphology the specimen most closely resembles Palaeortyx gallica which has been recorded from the late Oligocene to Middle Miocene of France and Germany.

The phylogenetic affinities of Palaeortyx are debated. Because of the presence of a marked, double fossa pneumotricipitalis, the taxon was originally considered to be a member of the Odontophorinae (Milne-Edwards 1867–1871).

However, a double fossa pneumotricipitalis also occurs in some stem group Galliformes (Mourer- Chauviré 1992, Mayr 2000) and Ammoperdix (Phasianinae) and appears to have evolved inde- pendently several times within Galliformes, unless it is plesiomorphic for galliform birds. As

evidenced by the Enspel galliform, Palaeortyx does not exhibit the derived beak morphology of Odontophorinae. Ballmann (1969) assumed a closer relationship between Palaeortyx and hill par- tridges of the genus Arborophila, but this hypothe- sis was based on overall similarity only. Although the pelvis of Palaeortyx agrees with that of Arborophila in the absence of well-developed tubercula praeacetabularia, the plesiomorphic limb proportions (femur subequal to humerus in length, as in Paleogene stem group Galliformes) rather support a position of Palaeortyx outside crown group “Phasianinae”. A definitive phyloge- netic assignment of Palaeortyx to any phasianid subclade has to await a better understanding of the relationships between the extant taxa. In any case, Mlíkovský's (2002) synonymization of Palaeortyx with the extant taxon Coturnix is incom- prehensible and without any foundation (see also Göhlich & Mourer-Chauviré 2005).

Extant galliform birds occur in a great variety of habitats, from semi-deserts to tropical forests and subantarctic tundra (del Hoyo et al. 1994), but little is known about the ecological preferences of Paleogene galliform birds. The palaeoenviron- ment surrounding Lake Enspel was dominated by a mesophytic forest, which probably reached close to the lake (Köhler 1998). Thus, the Enspel galli- form apparently was a forest-dwelling species.

The specimen is most remarkable for the preser- vation of a large accumulation of gizzard stones, which constitutes the earliest fossil evidence for grit ingestion in Phasianidae (the only other fossil record of gizzard stones in galliform birds comes from the stem group galliform Taubacrex Alva- renga, 1988 from the late Oligocene or early Miocene of Brazil; Alvarenga 1988, Mourer- Chauviré 2000). The fact that grit is found not only in the area of the gizzard but also in that of the crop is especially noteworthy. To the best of our knowledge, the accumulation of grit in the crop of extant birds has not been recorded so far, and the present specimen provides the first evidence of it in a fossil avian taxon.

In the phasianid galliforms, the gizzard stones mainly serve for grinding seeds and other coarse plant matter (Gionfriddo & Best 1999). On osteo- logical grounds it has been argued that early Eocene stem group Galliformes did not possess a large crop and fed on softer plant matter rather than hard-shelled seeds (Mayr 2006). Gizzard stones have not been reported for comparably well-preserved articulated skeletons of early Eocene stem group galliforms (Mayr 2000, 2006,

Mayr & Weidig 2004), but this negative evidence from few specimens does not conclusively prove their absence in vivo.

The Enspel Fossillagerstätte is well known for exceptional cases of soft tissue preservation, including the gliding membrane of an eomyid rodent (Storch et al. 1996, Engesser & Storch 1999).

Examination of preserved soft tissues in a tadpole and a coprolite showed that these consist of the fossilized remains of bacterial biofilms (Toporski et al. 2002). The skeleton of Palaeortyx likewise shows a halo of dark organic substance, which appears especially prominent in the neck and caudal body portion. This halo may mark the presence of down feathers whose original outlines became obscured. Few traces of contour feathers with pre- served outlines can be identified in the area of the right wing, including faint remains of some pri- maries. Bird carcasses tend to disarticulate rapidly, including loss of their body contour feathers (Davis & Briggs 1995). Thus, the Enspel Palaeortyx was probably buried soon after death, although we could not find an explanation for the fact that only few remains of the primaries are preserved.

The presence of relatively coarse sediment on the same bedding plane as the fossil suggests that the carcass fell into the lake in connection with a severe meteorological event, such as a storm or heavy rain that mobilized and transported coarse sediment grains from the shore into the deeper layers of the lake.

ACKNOWLEDGEMENTS

We thank S. Tränkner for taking the photo- graphs of the transferred specimen, and the refer- ees, C. Mourer-Chauviré and A. Elżanowski, for comments which improved the manuscript.

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STRESZCZENIE [Prawie kompletny szkielet Palaeortyx, kopal- nego grzebiącego z późnego Oligocenu Niemiec]

Kurowate (Phasianidae) nie występują w Eu- ropie przed końcem wczesnego oligocenu, a w całym paleogenie zachowały się głównie ich izolowane kości. Opisany tutaj jest prawie kom- pletny i zachowany w naturalnym zestawieniu szkielet kopalnego kurowatego z późnego oligo- cenu Enspel w Niemczech. Okaz ten zaliczony został do kurowatych na podstawie obecności processus intermetacarpalis kości lotkowej (car- pometacarpus), i do rodzaju Palaeortyx, do którego należą najliczniejsze kurowate oligocenu Europy, na podstawie proporcji kończyn, w szczególności równej długości kości udowej (femur) i kości ramiennej (humerus). Okaz ten po raz pierwszy pozwala na rozpoznanie szczegółów budowy czaszki i miednicy u późnooligoceńskiego kurowa- tego. Liczne gastrolity zachowały się zarówno w położeniu żołądka (mięśniowego) jak i wola.

Stanowi to najstarsze znalezisko gastrolitów u kurowatych.