A nearly complete skeleton of the fossil galliform bird ...2Landesamt für Denkmalpflege Rheinland-Pfalz, Referat Erdgeschichtliche Denkmalpflege, Große Langgasse 29, D-55116 Mainz,

INTRODUCTION

As traditionally recognized (e.g. Stresemann1927–1934), the galliform taxon Phasianidaeincludes 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 (47million years ago, Mayr 2000, 2006), all Eocene andearly Oligocene galliform taxa are outside crowngroup Galliformes and belong to the Gallinuloi-didae, Quercymegapodiidae, and Paraortygidae(Mourer-Chauviré 1992, Mayr 2005). Phasianidaedo not occur in Europe before the end of the earlyOligocene, about 30 million years ago (Mourer-Chauviré 1992), and most Oligocene phasianidsbelong to Palaeortyx Milne-Edwards, 1869. Thistaxon also occurs in the early Miocene of Franceand Germany (Göhlich & Mourer-Chauviré 2005)and has so far been known from isolated limb and

pectoral girdle bones only. The only otherphasianid from the Oligocene of Europe isSchaubortyx keltica (Eastman 1905), which is basedon a rather poorly preserved disarticulated skele-ton from the early Oligocene (MP 25) of France(Eastman 1905, Schaub 1945) and distinctly differsfrom Palaeortyx in its intermembral proportions(Mourer-Chauviré 1992; contra Mlíkovský 2002who synonymized Schaubortyx Brodkorb, 1964and 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; Mertzet al. in press) maar lake deposit of Enspel nearBad Marienberg in Westerwald, Germany. It is thesecond 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 millimetersand indicate the maximum length of the bone

A nearly complete skeleton of the fossil galliform bird Palaeortyx fromthe late Oligocene 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] 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 thelate 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 Paleogenefossil 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 Oligocenephasianid. 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

ACTA ORNITHOLOGICAVol. 41 (2006) No. 2

130 G. Mayr et al.

along its longitudinal axis. The specimen was pre-pared with the resin transfer method (Kühne1961).

SYSTEMATIC PALEONTOLOGY

Galliformes Temminck, 1820Phasianidae 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 fürDenkmalpflege Rheinland-Pfalz, Mainz (the spec-imen will be transferred to the Landessammlungfür 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.

Late Oligocene fossil galliform bird 131

as given by Göhlich & Mourer-Chauviré 2005).Skull, 48.7. Beak, length from nasofrontal hinge totip, ~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 wingand hindlimb elements of Palaeortyx were alreadydescribed in detail (Milne-Edwards 1867–1871,Ballmann 1969, Mourer-Chauviré 1992), and thusthe following description focuses on previouslyunknown skeletal elements.

The skull of Palaeortyx has so far been un-known and the Enspel galliform actually providesthe first skull details of an Oligocene phasianid(Fig. 3), the skull of Schaubortyx being very poorlypreserved. The beak measures about one third ofthe entire skull length and resembles that of theCommon 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 osnasale, which borders the caudal end of the largenarial opening (Fig. 3), is wide as in the Cracidae(guans, chachalacas, currasows) and Numidinae,wider than in most “Phasianinae” (except, forexample, Afropavo and Pavo). The os praefrontale iswell-developed and similar in shape to the corre-sponding bone of P. perdix; this bone is morereduced in, e.g., Coturnix, Rollulus, and Ammo-perdix, and articulates in a recess of the os frontalein Numidinae. As in other Phasianidae, the de-scending process of the os praefrontale is greatlyreduced and does not reach the jugal bar.Although the left sclerotic ring is completely pre-served, the exact number of ossicles cannot becounted but may have been 14 or 15 as in extantGalliformes. The processus postorbitalis is welldeveloped as in other Phasianidae (Fig. 3); thisprocess is very short in the Australasian Mega-podiidae (brush-turkeys, scrubfowl, megapodes)and the Gallinuloididae (Mayr & Weidig 2004).The processus zygomaticus, however, appears tohave been short, whereas it is very long and fuseswith the processus postorbitalis via an ossifiedaponeurosis zygomatica in Cracidae and mostPhasianidae; only in Numidinae it is vestigial. Asin other Phasianidae except for the short-billedOdontophorinae, the processus orbitalis of thequadratum is elongated and slender (Fig. 3). Themandible bears a well-developed processus retro-articularis; a fenestra mandibulae is absent (in

Fig. 2. Palaeortyx cf. gallica Milne-Edwards, 1869, articulatedskeleton from the late Oligocene of Enspel, Germany (PW2005/5023a-LS); photo taken before the fossil was transferredonto the artificial matrix. A dark halo representing fossilizedfeathers is clearly visible. Scale bar equals 10 mm.

contrast to, e.g., Tetraoninae in which it is verylarge). The well-preserved hyoid apparatus (Fig.3), which is also visible through the reverse of thetransparent slab, matches that of extant phasian-ids.

The morphology of the cervical vertebrae cor-responds to that of extant Phasianidae. As in thelatter, the third and fourth cervical vertebraeexhibit an osseous bridge from the processustransversus to the processus articularis caudalis(Fig. 3). Also as in extant Phasianidae, there are 20praesacral vertebrae, and there is one free thoracicvertebra 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 coracoidis shallow (PW 2005/5023b-LS) which is a de-rived characteristic of crown group Galliformes(Mourer-Chauviré 1992, Mayr 2000); the dorsalsurface 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 furculaand scapula prevents recognition of osteologicaldetails.

The sternum of Palaeortyx so far has not beendescribed and is visible from its right side in PW2005/5023a-LS. The carina sterni is well-devel-oped, with a pointed apex carinae and a convexcranial margin which bears a marked sulcus cari-nae. The incisions in the caudal margin are verydeep as in extant Odontophorinae and “Phasia-ninae”, although it is difficult to discern the tra-beculae (the trabecula intermedia extends along acrack in the original slab, the caudal portion of thetrabecula lateralis overlays the cranial end of thesynsacrum). Details of the cranial portion of thisbone, including the shape of the spina externa, arenot visible.

The caudal surface of the proximal humerus ofPalaeortyx 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 caudalsurface of the proximal humerus was exposedbefore embedding in the resin and can still beseen through the reverse of the transparent slab.The carpometacarpus exhibits a processus inter-metacarpalis (Fig. 4) which is a derived feature ofthe Phasianidae and absent in Paleogene stemgroup Galliformes; this process is secondarilyreduced in Numidinae (Stegmann 1978). As inmany extant Phasianidae (e.g. Alectoris chukar),there is an ossified tendon along the craniodorsalsurface of the carpometacarpus. The phalanxproximalis digiti majoris is not as abbreviated andcraniocaudally wide as in some extant “Phasia-ninae” (e.g., Rollulus and Lophura). The phalanxdigiti alulae bears a small ungual phalanx which iscomparable in size to that of Alectoris.

The pelvis appears to have been of similar pro-portions to that of most extant Phasianidae andnot as wide as in the Tetraoninae, although itsoriginal proportions are difficult to reconstruct

Late Oligocene fossil galliform bird 133

because of to the flattening of the specimen. Mostnotably, the tubercula praeacetabularia are verysmall as in extant Odontophorinae (Holman1964), Tetraoninae, and a few taxa within “Phasia-ninae”, such as Ammoperdix and, according toDyke et al. (2003), Arborophila and Galloperdix (Fig.4); they are much more developed in otherPhasianidae.

As in Palaeortyx, the femur is subequal to thehumerus in length, whereas it is much longer inSchaubortyx keltica, extant “Phasianinae”, and mostOdontophorinae. The femur is shorter than orsubequal to the humerus in stem group Gal-liformes and extant Cracidae, Numidinae, Cyr-tonyx (Odontophorinae; Holman 1961), andTetraoninae. Only the distal end of the tibiotarsusshows osteological details and resembles the distaltibiotarsus of Palaeortyx. The tarsometatarsus alsoagrees with that of Palaeortyx in its proportionsand morphology. It lacks a spur which withinextant Phasianidae is also wanting in the males ofOdontophorinae, Numidinae, and Tetraoninae(contra Dyke et al. 2003), as well as Perdix (contraDyke et al. 2003), Margaroperdix, Melanoperdix,Coturnix, Ammoperdix, Rollulus, Arborophila (contraDyke 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 veryshort. An ossified tendon attaches to the processusarticularis tarsometatarsalis of the os metatarsale I and extends along the plantar surface of the tarsometatarsus.

Although remains of the primaries of the rightwing are preserved, they do not permit a mean-ingful description. Ornamental feathers (e.g., onthe 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 feweramount of grit is also visible in the crop area, nextto 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 arewhite and grayish quartz pebbles, about one fifthis reddish pyroclastic material whereas almost all grit particles in the crop area are reddish. A brown organic substance with a fibrous texturesuggestive of plant material is preserved in some places between the gizzard stones. Someunidentifiable remains of seeds are preserved inthe crop area.

Fig. 4. Palaeortyx cf. gallica Milne-Edwards, 1869 (PW2005/5023a-LS); distal part of the left wing (A) and pelvis withgizzard stones (B). Abbreviations: pim — processus inter-metacarpalis, tpa — tuberculum praeacetabulare. Specimencoated with ammonium chloride. Scale bars equal 5 mm.

DISCUSSION

The derived presence of a processus inter-metacarpalis on the carpometacarpus supportsclassification of the Enspel galliform into Phasia-nidae. It is assigned to Palaeortyx, the most abun-dant phasianid taxon in the late Oligocene ofEurope, because of its intermembral proportionswith the femur being as long as the humerus. Thepreservation does not permit detailed osteologicalcomparisons, but in size and morphology thespecimen most closely resembles Palaeortyx gallicawhich has been recorded from the late Oligoceneto Middle Miocene of France and Germany.

The phylogenetic affinities of Palaeortyx aredebated. Because of the presence of a marked,double fossa pneumotricipitalis, the taxon wasoriginally considered to be a member of theOdontophorinae (Milne-Edwards 1867–1871).However, a double fossa pneumotricipitalis alsooccurs 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 doesnot exhibit the derived beak morphology ofOdontophorinae. Ballmann (1969) assumed acloser relationship between Palaeortyx and hill par-tridges of the genus Arborophila, but this hypothe-sis was based on overall similarity only. Althoughthe pelvis of Palaeortyx agrees with that ofArborophila in the absence of well-developedtubercula praeacetabularia, the plesiomorphiclimb proportions (femur subequal to humerus inlength, as in Paleogene stem group Galliformes)rather support a position of Palaeortyx outsidecrown group “Phasianinae”. A definitive phyloge-netic assignment of Palaeortyx to any phasianidsubclade has to await a better understanding ofthe relationships between the extant taxa. In anycase, Mlíkovský's (2002) synonymization ofPalaeortyx with the extant taxon Coturnix is incom-prehensible and without any foundation (see alsoGöhlich & Mourer-Chauviré 2005).

Extant galliform birds occur in a great varietyof habitats, from semi-deserts to tropical forestsand subantarctic tundra (del Hoyo et al. 1994), butlittle is known about the ecological preferences ofPaleogene galliform birds. The palaeoenviron-ment surrounding Lake Enspel was dominated bya mesophytic forest, which probably reached closeto 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 forgrit ingestion in Phasianidae (the only other fossilrecord of gizzard stones in galliform birds comesfrom the stem group galliform Taubacrex Alva-renga, 1988 from the late Oligocene or earlyMiocene of Brazil; Alvarenga 1988, Mourer-Chauviré 2000). The fact that grit is found not onlyin the area of the gizzard but also in that of thecrop is especially noteworthy. To the best of ourknowledge, the accumulation of grit in the crop ofextant birds has not been recorded so far, and thepresent specimen provides the first evidence of itin a fossil avian taxon.

In the phasianid galliforms, the gizzard stonesmainly serve for grinding seeds and other coarseplant matter (Gionfriddo & Best 1999). On osteo-logical grounds it has been argued that earlyEocene stem group Galliformes did not possess alarge crop and fed on softer plant matter ratherthan hard-shelled seeds (Mayr 2006). Gizzardstones have not been reported for comparablywell-preserved articulated skeletons of earlyEocene stem group galliforms (Mayr 2000, 2006,

Mayr & Weidig 2004), but this negative evidencefrom few specimens does not conclusively provetheir absence in vivo.

The Enspel Fossillagerstätte is well known forexceptional cases of soft tissue preservation,including the gliding membrane of an eomyidrodent (Storch et al. 1996, Engesser & Storch 1999).Examination of preserved soft tissues in a tadpoleand a coprolite showed that these consist of thefossilized remains of bacterial biofilms (Toporski etal. 2002). The skeleton of Palaeortyx likewise showsa halo of dark organic substance, which appearsespecially prominent in the neck and caudal bodyportion. This halo may mark the presence ofdown feathers whose original outlines becameobscured. Few traces of contour feathers with pre-served outlines can be identified in the area of theright 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 Palaeortyxwas probably buried soon after death, althoughwe could not find an explanation for the fact thatonly few remains of the primaries are preserved.The presence of relatively coarse sediment on thesame bedding plane as the fossil suggests that thecarcass fell into the lake in connection with asevere meteorological event, such as a storm orheavy rain that mobilized and transported coarsesediment grains from the shore into the deeperlayers 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, forcomments 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 ichizolowane kości. Opisany tutaj jest prawie kom-pletny i zachowany w naturalnym zestawieniuszkielet kopalnego kurowatego z późnego oligo-cenu Enspel w Niemczech. Okaz ten zaliczonyzostał do kurowatych na podstawie obecnościprocessus intermetacarpalis kości lotkowej (car-pometacarpus), i do rodzaju Palaeortyx, do któregonależą najliczniejsze kurowate oligocenu Europy,na podstawie proporcji kończyn, w szczególnościrównej długości kości udowej (femur) i kościramiennej (humerus). Okaz ten po raz pierwszypozwala na rozpoznanie szczegółów budowyczaszki i miednicy u późnooligoceńskiego kurowa-tego. Liczne gastrolity zachowały się zarówno wpołożeniu żołądka (mięśniowego) jak i wola.Stanowi to najstarsze znalezisko gastrolitów ukurowatych.

Late Oligocene fossil galliform bird 135

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