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Home , Ammoperdix, Galliformes, Galloperdix, Meleagridinae, Palaeortyx, Partridge

ACTA ORNITHOLOGICA Vol. 41 (2006) No. 2

A nearly complete skeleton of the galliform from the late of Germany

Gerald MAYR1, Markus POSCHMANN2 & Michael WUTTKE2

1Forschungsinstitut Senckenberg, Sektion für Ornithologie, Senckenberganlage 25, D-60325 Frankfurt am Main, GERMANY, e-mail: [email protected] 2Landesamt für Denkmalpflege Rheinland-Pfalz, Referat Erdgeschichtliche Denkmalpflege, Große Langgasse 29, D-55116 Mainz, GERMANY, e-mail: [email protected]

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 do not occur in before the end of the early Oligocene, and their fossil record mainly comprises isolated . 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, , Palaeortyx, gizzard stones, Oligocene

Received — July 2006, accepted — Oct. 2006

INTRODUCTION pectoral girdle bones only. The only other phasianid from the Oligocene of Europe is As traditionally recognized (e.g. Stresemann Schaubortyx keltica (Eastman 1905), which is based 1927–1934), the galliform on a rather poorly preserved disarticulated skele- includes the Odontophorinae ( ), ton from the early Oligocene (MP 25) of Numidinae (), (turkeys), (Eastman 1905, Schaub 1945) and distinctly differs Tetraoninae () and the, probably para- from Palaeortyx in its intermembral proportions phyletic, “” ( quails, pheas- (Mourer-Chauviré 1992; contra Mlíkovský 2002 , , etc.; Kimball et al. 1999, Arm- who synonymized Schaubortyx Brodkorb, 1964 strong et al. 2001). Of these, only Tetraoninae and and Palaeortyx Milne-Edwards, 1869). “Phasianinae” occur in Europe today. Here we describe a nearly complete and artic- Although the European fossil record of galli- ulated skeleton of Palaeortyx from the late Oligo- form birds dates back into the early (47 cene (MP 28, Storch et al. 1996; i.e. 24.7 mya; Mertz million ago, Mayr 2000, 2006), all Eocene and et al. in press) maar lake deposit of Enspel near early Oligocene galliform taxa are outside crown Bad Marienberg in Westerwald, Germany. It is the group Galliformes and belong to the Gallinuloi- second avian specimen from Enspel (Mayr 2001), didae, Quercymegapodiidae, and Paraortygidae and one of a few articulated skeletons of Oligo- (Mourer-Chauviré 1992, Mayr 2005). Phasianidae cene galliform birds. do not occur in Europe before the end of the early Oligocene, about 30 million years ago (Mourer- Chauviré 1992), and most Oligocene phasianids MATERIAL AND METHODS belong to Palaeortyx Milne-Edwards, 1869. This taxon also occurs in the early of France Osteological terminology follows Baumel & and Germany (Göhlich & Mourer-Chauviré 2005) Witmer (1993). Measurements are in millimeters and has so far been known from isolated limb and and indicate the maximum length of the 130 G. Mayr et al. along its longitudinal axis. The specimen was pre- Referred specimen. PW 2005/5023a-LS (Figs. 1–4) pared with the resin transfer method (Kühne and PW 2005/5023b-LS (few bones only, not fig- 1961). ured), currently deposited in Landesamt für Denkmalpflege Rheinland-Pfalz, Mainz (the spec- imen will be transferred to the Landessammlung SYSTEMATIC PALEONTOLOGY für Naturkunde RLP, Germany). Locality and horizon. Enspel near Bad Marien- Galliformes Temminck, 1820 berg in Westerwald, Rheinland-Pfalz, Germany; Phasianidae sensu Stresemann (1927–34) late Oligocene, MP 28 (Storch et al. 1996). Palaeortyx cf. gallica Milne-Edwards, 1869 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 , lfe — left , lhu — left humerus, ltb — left tibiotarsus, ltm — left , 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. Late Oligocene fossil galliform bird 131 as given by Göhlich & Mourer-Chauviré 2005). Skull, 48.7. , 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 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 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 (guans, , currasows) and Numidinae, wider than in most “Phasianinae” (except, for Fig. 2. Palaeortyx cf. gallica Milne-Edwards, 1869, articulated example, Afropavo and ). The os praefrontale is skeleton from the late Oligocene of Enspel, Germany (PW well-developed and similar in shape to the corre- 2005/5023a-LS); photo taken before the fossil was transferred sponding bone of P. perdix; this bone is more onto the artificial matrix. A dark halo representing fossilized reduced in, e.g., , Rollulus, and Ammo- is clearly visible. Scale bar equals 10 mm. 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 contrast to, e.g., Tetraoninae in which it is very reduced and does not reach the jugal bar. large). The well-preserved hyoid apparatus (Fig. Although the left sclerotic ring is completely pre- 3), which is also visible through the reverse of the served, the exact number of ossicles cannot be transparent slab, matches that of extant phasian- counted but may have been 14 or 15 as in extant ids. Galliformes. The processus postorbitalis is well The morphology of the cervical vertebrae cor- developed as in other Phasianidae (Fig. 3); this responds to that of extant Phasianidae. As in the process is very short in the Australasian Mega- latter, the third and fourth cervical vertebrae podiidae (brush-turkeys, , ) exhibit an osseous bridge from the processus and the Gallinuloididae (Mayr & Weidig 2004). transversus to the processus articularis caudalis The processus zygomaticus, however, appears to (Fig. 3). Also as in extant Phasianidae, there are 20 have been short, whereas it is very long and fuses praesacral vertebrae, and there is one free thoracic with the processus postorbitalis via an ossified vertebra between the synsacrum and the notari- aponeurosis zygomatica in Cracidae and most um (part of which can be seen in PW 2005/5023b- Phasianidae; only in Numidinae it is vestigial. As LS). The tail vertebrae are not preserved. in other Phasianidae except for the short-billed The facies articularis scapularis of the coracoid Odontophorinae, the processus orbitalis of the is shallow (PW 2005/5023b-LS) which is a de- quadratum is elongated and slender (Fig. 3). The rived characteristic of Galliformes mandible bears a well-developed processus retro- (Mourer-Chauviré 1992, Mayr 2000); the dorsal articularis; a fenestra mandibulae is absent (in surface of the extremitas sternalis does not exhibit 132 G. Mayr et al.

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 the Enspel galliform, although a part of the caudal and scapula prevents recognition of osteological surface of the proximal humerus was exposed details. before embedding in the resin and can still be The sternum of Palaeortyx so far has not been seen through the reverse of the transparent slab. described and is visible from its right side in PW The carpometacarpus exhibits a processus inter- 2005/5023a-LS. The carina sterni is well-devel- metacarpalis (Fig. 4) which is a derived feature of oped, with a pointed apex carinae and a convex the Phasianidae and absent in Paleogene stem cranial margin which bears a marked sulcus cari- group Galliformes; this process is secondarily nae. The incisions in the caudal margin are very reduced in Numidinae (Stegmann 1978). As in deep as in extant Odontophorinae and “Phasia- many extant Phasianidae (e.g. chukar), ninae”, although it is difficult to discern the tra- there is an ossified tendon along the craniodorsal beculae (the trabecula intermedia extends along a surface of the carpometacarpus. The phalanx crack in the original slab, the caudal portion of the proximalis digiti majoris is not as abbreviated and trabecula lateralis overlays the cranial end of the craniocaudally wide as in some extant “Phasia- synsacrum). Details of the cranial portion of this ninae” (e.g., Rollulus and Lophura). The phalanx bone, including the shape of the spina externa, are digiti alulae bears a small ungual phalanx which is not visible. comparable in size to that of Alectoris. The caudal surface of the proximal humerus of The pelvis appears to have been of similar pro- Palaeortyx is very characteristic owing to the pres- portions to that of most extant Phasianidae and ence of a marked, double fossa pneumotricipitalis. not as wide as in the Tetraoninae, although its Unfortunately, this feature is not clearly visible in original proportions are difficult to reconstruct Late Oligocene fossil galliform bird 133 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 and, according to Dyke et al. (2003), and (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 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 Fig. 4. Palaeortyx cf. gallica Milne-Edwards, 1869 (PW Dyke et al. 2003), Margaroperdix, Melanoperdix, 2005/5023a-LS); distal part of the left wing (A) and pelvis with Coturnix, Ammoperdix, Rollulus, Arborophila (contra gizzard stones (B). Abbreviations: pim — processus inter- metacarpalis, tpa — tuberculum praeacetabulare. Specimen Dyke et al. 2003), , and Ptilopachus. coated with ammonium chloride. Scale bars equal 5 mm. 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 DISCUSSION articularis tarsometatarsalis of the os metatarsale I and extends along the plantar surface of the The derived presence of a processus inter- tarsometatarsus. metacarpalis on the carpometacarpus supports Although remains of the primaries of the right classification of the Enspel galliform into Phasia- wing are preserved, they do not permit a mean- nidae. It is assigned to Palaeortyx, the most abun- ingful description. Ornamental feathers (e.g., on dant phasianid taxon in the late Oligocene of the head) cannot be discerned. Europe, because of its intermembral proportions A large quantity of grit is preserved in a well- with the femur being as long as the humerus. The defined area of about 21 x 28.5 mm, between preservation does not permit detailed osteological the pelvis and the sternum, and unquestionably comparisons, but in size and morphology the represents gizzard stones (Figs. 1 and 4). A fewer specimen most closely resembles Palaeortyx gallica amount of grit is also visible in the crop area, next which has been recorded from the late Oligocene to the proximal end of the right humerus (Fig. 1). to Middle Miocene of France and Germany. Grit size varies between 0.1 and 2.6 mm, on The phylogenetic affinities of Palaeortyx are average the particles have a diameter of 1.5 mm; debated. Because of the presence of a marked, all are rounded. Most of the gizzard stones are double fossa pneumotricipitalis, the taxon was white and grayish quartz pebbles, about one fifth originally considered to be a member of the is reddish pyroclastic material whereas almost Odontophorinae (Milne-Edwards 1867–1871). all grit particles in the crop area are reddish. However, a double fossa pneumotricipitalis also A brown organic substance with a fibrous texture occurs in some stem group Galliformes (Mourer- suggestive of plant material is preserved in Chauviré 1992, Mayr 2000) and Ammoperdix some places between the gizzard stones. Some (Phasianinae) and appears to have evolved inde- unidentifiable remains of seeds are preserved in pendently several times within Galliformes, the crop area. unless it is plesiomorphic for galliform birds. As 134 G. Mayr et al. evidenced by the Enspel galliform, Palaeortyx does Mayr & Weidig 2004), but this negative evidence not exhibit the derived beak morphology of from few specimens does not conclusively prove Odontophorinae. Ballmann (1969) assumed a their absence in vivo. closer relationship between Palaeortyx and hill par- The Enspel Fossillagerstätte is well known for tridges of the Arborophila, but this hypothe- exceptional cases of soft tissue preservation, sis was based on overall similarity only. Although including the gliding membrane of an eomyid the pelvis of Palaeortyx agrees with that of (Storch et al. 1996, Engesser & Storch 1999). Arborophila in the absence of well-developed Examination of preserved soft tissues in a tadpole tubercula praeacetabularia, the plesiomorphic and a coprolite showed that these consist of the limb proportions (femur subequal to humerus in fossilized remains of bacterial biofilms (Toporski et length, as in Paleogene stem group Galliformes) al. 2002). The skeleton of Palaeortyx likewise shows rather support a position of Palaeortyx outside a halo of dark organic substance, which appears crown group “Phasianinae”. A definitive phyloge- especially prominent in the neck and caudal body netic assignment of Palaeortyx to any phasianid portion. This halo may mark the presence of subclade has to await a better understanding of down feathers whose original outlines became the relationships between the extant taxa. In any obscured. Few traces of contour feathers with pre- case, Mlíkovský's (2002) synonymization of served outlines can be identified in the area of the Palaeortyx with the extant taxon Coturnix is incom- right wing, including faint remains of some pri- prehensible and without any foundation (see also maries. Bird carcasses tend to disarticulate rapidly, Göhlich & Mourer-Chauviré 2005). including loss of their body contour feathers Extant galliform birds occur in a great variety (Davis & Briggs 1995). Thus, the Enspel Palaeortyx of habitats, from semi- to tropical forests was probably buried soon after death, although and subantarctic tundra (del Hoyo et al. 1994), but we could not find an explanation for the fact that little is known about the ecological preferences of only few remains of the primaries are preserved. Paleogene galliform birds. The palaeoenviron- The presence of relatively coarse sediment on the ment surrounding Lake Enspel was dominated by same bedding plane as the fossil suggests that the a mesophytic forest, which probably reached close carcass fell into the lake in connection with a to the lake (Köhler 1998). Thus, the Enspel galli- severe meteorological event, such as a storm or form apparently was a forest-dwelling . heavy rain that mobilized and transported coarse The specimen is most remarkable for the preser- sediment grains from the shore into the deeper vation of a large accumulation of gizzard stones, layers of the lake. which constitutes the earliest fossil evidence for grit ingestion in Phasianidae (the only other fossil record of gizzard stones in galliform birds comes ACKNOWLEDGEMENTS from the stem group galliform Taubacrex Alva- renga, 1988 from the late Oligocene or early We thank S. Tränkner for taking the photo- Miocene of Brazil; Alvarenga 1988, Mourer- graphs of the transferred specimen, and the refer- Chauviré 2000). The fact that grit is found not only ees, C. Mourer-Chauviré and A. Elżanowski, for in the area of the gizzard but also in that of the comments which improved the manuscript. 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 REFERENCES present specimen provides the first evidence of it in a fossil avian taxon. Alvarenga H. 1988. Aves Fóssil (: Rallidae) dos Folhelhos da Bacia de Taubaté, Estado de Sno Paulo, Brasil. In the phasianid galliforms, the gizzard stones Anais da Academia Brasileira de CiLncias 60: 321–328. mainly serve for grinding seeds and other coarse Armstrong M. H., Braun E. L., Kimball R. T. 2001. Phylogenetic plant matter (Gionfriddo & Best 1999). 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