Post on 08-Aug-2020
RESEARCH PAPER
New specimens of the avian taxa Eurotrochilus (Trochilidae)and Palaeotodus (Todidae) from the early Oligocene of Germany
Gerald Mayr • Norbert Micklich
Received: 8 September 2009 / Accepted: 15 December 2009 / Published online: 9 January 2010
� Springer-Verlag 2010
Abstract We describe new specimens of stem group
representatives of Trochilidae (hummingbirds) and Todi-
dae (todies) from the Rupelian of Frauenweiler in southern
Germany. The hummingbird fossil constitutes the fourth
record of Eurotrochilus inexpectatus. It consists only of
wing and pectoral girdle elements, but shows the previ-
ously unknown crista deltopectoralis of the humerus,
whose shape differs from modern hummingbirds. The
carpometacarpus bears a well-developed processus inter-
metacarpalis, which is a further synapomorphy of Eurot-
rochilus and crown group Trochilidae. The disarticulated
partial skeleton of the tody allows a definitive taxonomic
assignment of the Frauenweiler species to Palaeotodus
itardiensis Mourer-Chauvire, and likewise exhibits so far
unknown osteological details, including the morphologies
of the quadrate and scapula. We further comment on the
exceptional taphonomy and preservation of avian fossils
from the Frauenweiler clay pit, where terrestrial birds are
represented only by small to very small species, whose
skeletons are always strongly disarticulated.
Keywords Aves � Fossil birds � Rupelian � Frauenweiler �Taphonomy
Kurzfassung Wir beschreiben neue Exemplare von
Stammgruppenvertretern der Trochilidae (Kolibris) und
Todidae (Todis) aus dem Rupelton von Frauenweiler in
Suddeutschland. Das Kolibrifossil ist der vierte Nachweis
von Eurotrochilus inexpectatus. Es besteht nur aus Flugel-
und Brustgurtelelementen, zeigt aber die bisher unbekannte
Crista deltopectoralis des Humerus, deren Form sich von
derjenigen moderner Kolibris unterscheidet. Der gut ent-
wickelte Processus intermetacarpalis des Carpometacarpus
stellt eine weitere Synapomorphie von Eurotrochilus
und Kronengruppen-Trochilidae dar. Das disartikulierte
Teilskelett des Todi erlaubt eine endgultige taxonomische
Zuordnung der Frauenweiler-Art zu Palaeotodus itardien-
sis Mourer-Chauvire und weist ebenfalls bis jetzt unbe-
kannte osteologische Details auf, einschließlich der
Morphologien des Quadratums und der Scapula. Wir
diskutieren daruber hinaus die ungewohnliche Taphonomie
und der Erhaltung der Vogelfossilien aus der Tongrube
Frauenweiler, in welcher terrestrische Vogel nur durch
kleine bis sehr kleine Arten reprasentiert sind, deren
Skelette zudem immer stark zerfallen sind.
Schlusselworter Aves � fossile Vogel � Rupelium �Frauenweiler � Taphonomie
Introduction
The early Oligocene marine sediments of the Frauenweiler
clay pit near Heidelberg in southern Germany were
deposited during a transgression of the Rupelian sea, and
have yielded a rich ichthyofauna (Micklich and Parin 1996;
Micklich 1998). Bird fossils were occasionally found
by amateur collectors searching for fishes, but their study
has only begun in the past decade. The locality is notable
for the preservation of very small avian species, and
well-preserved remains of new or poorly known taxa
were discovered. Aquatic birds are represented by the
G. Mayr (&)
Forschungsinstitut Senckenberg, Sektion Ornithologie,
Senckenberganlage 25, 60325 Frankfurt a.M., Germany
e-mail: Gerald.Mayr@senckenberg.de
N. Micklich
Natural History Department, Hessisches Landesmuseum
Darmstadt, Friedensplatz 1, 64283 Darmstadt, Germany
123
Palaontol Z (2010) 84:387–395
DOI 10.1007/s12542-009-0047-z
procellariiform Diomedeoididae (Mayr et al. 2002; Mayr
2009a) and the Gaviidae (loons; Mayr 2004a). Because the
Frauenweiler fossil site is close to the former shoreline
(Micklich and Hildebrandt 2005), however, many species
belong to terrestrial taxa. Among these are Turnicidae
(buttonquails; Mayr and Knopf 2007a), Coliiformes
(mousebirds; Mayr 2000), Trogoniformes (trogons; Mayr
2005a), Piciformes (woodpeckers and allies; Mayr 2005b,
2006), and Passeriformes (passerines; Mayr and Manegold
2004, 2006).
From a biogeographic point of view, certainly the most
interesting Frauenweiler birds are stem group representa-
tives of the Trochilidae (hummingbirds) and Todidae
(todies). Extant todies occur on the Greater Antilles and the
distribution of hummingbirds is today restricted to the New
World, so that modern todies and hummingbirds only
coexist in the Caribbean region. The Frauenweiler Troc-
hilidae belong to the species Eurotrochilus inexpectatus
Mayr, 2004b and were so far known from three partial
skeletons (Mayr 2004b, 2007). The tody was identified as
Palaeotodus cf. itardiensis Mourer-Chauvire, 1985 and
was previously represented by a single specimen (Mayr and
Knopf 2007b). Recent excavation campaigns in summer
2009 yielded new specimens of both species, which exhibit
previously unrecognized osteological features, and which
are described in the present study.
Materials and methods
Osteological terminology follows Baumel and Witmer
(1993); the description focuses on features not already
mentioned by Mayr (2004b, 2007) and Mayr and Knopf
(2007b). Institutional abbreviations: HLMD—Hessisches
Landesmuseum, Darmstadt, Germany; SMNK—Staatli-
ches Museum fur Naturkunde, Karlsruhe, Germany.
Systematic paleontology
• Apodiformes Peters, 1940
• Trochilidae Vigors, 1825
• Eurotrochilus Mayr, 2004b
• Eurotrochilus inexpectatus Mayr, 2004b
Referred specimen
SMNK-PAL.6599 (both wings and pectoral girdle ele-
ments, as well as a few ribs; part and counter part of the
specimen are combined on a single slab; Fig. 1). Found in
August 2009 by Harald and Annette Oechsler.
Locality and horizon
Frauenweiler south of Wiesloch (Baden-Wurttemberg,
Germany), former clay pit of the Bott-Eder GmbH (‘‘Grube
Unterfeld’’); Rupelian, early Oligocene (Micklich and
Hildebrandt 2005); layer 8 according to current profilings
(L. Hildebrandt, personal communication).
Measurements (length in mm)
Coracoid, 7.9 (left), 7.8 (right). Humerus, 6.4 (left), 6.4
(right). Ulna, 8.5 (left). Carpometacarpus, 6.8 (left), 6.9
(right).
Remarks
The taxon Eurotrochilus was first recognized in
Frauenweiler, but has recently also been discovered in
early Oligocene localities in Poland (Bochenski and
Bochenski 2008) and France (Louchart et al. 2008).
Although the osteology of Eurotrochilus is thus well
known, the new specimen shows some previously unknown
osteological features that justify its description.
Description and comparison
Both coracoids are preserved, and the shape of the bone is
for the first time completely visible. As in the more
primitive stem group hummingbird Jungornis tesselatus
Karhu, 1988 (Karhu 1999: Fig. 3), the medial margin is
markedly concave in the midsection of the bone. Other
morphological details correspond to previous descriptions
(Mayr 2004b, 2007). Likewise, both scapulae are visible in
the fossil. The extremitas cranialis exhibits a less derived
morphology than in crown group Trochilidae, in which the
straight end runs perpendicular to the long axis of the bone;
except for the proportionally smaller acromion, it more
closely resembles the extremitas cranialis of Jungornis
(Mayr 2007). The right extremitas omalis of the furcula is
situated next to the left carpometacarpus and exhibits a
distinct, circular facies articularis acrocoracoidea (Fig. 2h).
Humerus morphology of Eurotrochilus is comparatively
well known, but the crista deltopectoralis is not preserved
in the previous specimens (Mayr 2004b, 2007). This crest
is more protruding and pointed than in crown group
Trochilidae (Fig. 2), and resembles the crista deltopecto-
ralis of the less derived stem group hummingbirds Argornis
and Jungornis (Karhu 1988, 1999). The ulnae are well
preserved, but do not exhibit new osteological features.
The very large tuberculum carpale (Fig. 2f) is a further
synapomorphy of Eurotrochilus and crown group Trochi-
lidae, although this tubercle is still somewhat smaller and
388 G. Mayr, N. Micklich
123
less cranially protruding than in modern hummingbirds. As
in the latter, the incisura tendinosa is very marked.
The carpometacarpus bears a well-developed processus
intermetacarpalis, which is, however, slightly smaller than
in crown group Trochilidae (Fig. 2g–i). A small processus
intermetacarpalis was recognized in previous Eurotrochilus
specimens (Mayr 2004b), but in none of these is it as well
developed and clearly visible as in the new fossil. This
feature, which is absent in other apodiform birds, may
constitute another synapomorphy of Eurotrochilus and
crown group Trochilidae. According to Bochenski and
Bochenski (2008) a processus intermetacarpalis is absent in
Eurotrochilus noniewiczi Bochenski and Bochenski, 2008,
but the carpometacarpi of the only known specimen of this
species are rather poorly preserved, so that the alleged
absence of this feature may be an artifact of preservation.
The well-preserved distal end of the right carpometacarpus
confirms previous observations (Mayr 2004b), that contrary
to crown group Trochilidae, the os metacarpale minus of
Eurotrochilus does not protrude beyond the os metacarpale
majus. The caudal rim of the dorsal portion of the trochlea
carpalis is further more rounded that in crown group
Trochilidae. As in the latter, however, there is a narrow,
oblique fossa between the fossa supratrochlearis and the
processus intermetacarpalis (Fig. 2g).
• Alcediniformes sensu Mayr (1998)
• Todidae Vigors, 1825
• Palaeotodus Olson, 1976
• Palaeotodus itardiensis Mourer-Chauvire, 1985
Fig. 1 Eurotrochilus inexpectatus Mayr, 2004b from the early
Oligocene of Frauenweiler. Specimen SMNK-PAL.6599 (coated with
ammonium chloride) and interpretative drawings. Abbreviations: furfurcula, lcm left carpometacarpus, lco left coracoid, lhu left humerus,
lsc left scapula, lul left ulna, min phalanx digiti minoris, ocr right os
carpi radiale, ocu right os carpi ulnare, pdm phalanx distalis digiti
majoris, rad radius, rcm right carpometacarpus, rco right coracoid,
rhu right humerus, rsc right scapula, rul right ulna
New specimens of Eurotrochilus and Palaeotodus 389
123
Referred specimen
HLMD-WT 590 (disarticulated partial postcranial skeleton
on a slab; Fig. 3). Found in July 2009 during an excavation
campaign of HLMD.
Locality and horizon
Frauenweiler south of Wiesloch (Baden-Wurttemberg,
Germany), former clay pit of the Bott-Eder GmbH (‘‘Grube
Unterfeld’’); Rupelian, early Oligocene (Micklich and
Hildebrandt 2005); layer 9 according to current profilings
(L. Hildebrandt, personal communication).
Measurements (length in mm)
Coracoid, 15.1 (right). Humerus, 18.2 (left), *18.2 (right).
Ulna, 23.2 (left). Femur, 14.7 (left).
Remarks
Stem group representatives of todies have been found in
late Eocene and early Oligocene fossil sites of the Northern
Hemisphere (Mayr 2009b). All were classified in the taxon
Palaeotodus, which was originally erected for Palaeotodus
emryi Olson, 1976 from the early Oligocene of Wyoming
(Olson 1976). This species is known from the holotype
skull and a referred proximal humerus (Olson 1976), as
well as an undescribed postcranial skeleton (Mayr and
Knopf 2007b). Two species of Palaeotodus, P. itardiensis
Mourer-Chauvire, 1985 and P. escampsiensis Mourer-
Chauvire, 1985 were further described from the late
Eocene and early Oligocene of the Quercy fissure fillings in
France (Mourer-Chauvire 1985). Both are represented by
few bones: the holotype and only known specimen of the
late Eocene P. escampsiensis is an incomplete humerus; the
material assigned to the early Oligocene P. itardiensis
Fig. 2 Selected bones of Eurotrochilus inexpectatus Mayr, 2004b
from the early Oligocene of Frauenweiler (SMNK-PAL.6599). a–dright (a, c) and left (b, d) humerus in cranial (a, b) and caudal (c, d)
view, in comparison to e the left humerus of the Great-billed Hermit,
Phaethornis malaris (Trochilidae); f right ulna in craniodorsal view;
g right carpometacarpus in dorsal view, left ulna in dorsal view, and
distal left carpometacarpus in ventral view; h left carpometacarpus in
dorsal view, left ulna in cranioventral view, and right carpometacar-
pus in ventral view; i right carpometacarpus of Hairy Hermit, Glaucis
hirsuta (Trochilidae); j right coracoid in dorsal view, right scapula in
lateral view, left scapula in medial view, and left coracoid in ventralview. Abbreviations: cdp crista deltopectoralis, ext processus
extensorius, faa facies articularis acrocoracoidea, fos oblique fossa
proximal of processus intermetacarpalis, imc processus intermeta-
carpalis, lcm left carpometacarpus, lco left coracoid, lsc left scapula,
lul left ulna, rcm right carpometacarpus, rco right coracoid, rsc right
scapula, tbc tuberculum carpale. Fossil bones are coated with
ammonium chloride. Scale bars equal 2 mm
390 G. Mayr, N. Micklich
123
consists of the holotype, a proximal ulna, as well as a
referred distal tibiotarsus and proximal tarsometatarsus.
Mayr and Knopf (2007b) tentatively identified a disar-
ticulated postcranial skeleton from Frauenweiler as Pal-
aeotodus cf. itardiensis, but preservation did not allow for
detailed comparisons with the Quercy material. The ulna of
the new fossil can be directly compared with the holotype
of Palaeotodus itardiensis, so that a definitive assignment
of the Frauenweiler tody to that species is now possible.
Description and comparison
Of the skull elements, only the quadrates can be unam-
biguously identified, even though an elongate, flattened
bone may represent a portion of one of the mandibular rami
(Fig. 3). Both quadrates are visible in lateral view; the left
one lacks the processus orbitalis, whereas the more com-
plete right quadrate is poorly preserved (Fig. 4a, b). The
capitulum squamosum is separated from the rest of
the processus oticus by a narrow constriction (Fig. 4a). The
condylus medialis of the processus mandibularis has a
similar shape to that of extant Todidae and is somewhat
wider and less ventrally protruding than in Momotidae
(Fig. 4c); in contrast to extant Todidae but as in other
alcediniform birds, the lateral surface of its base is con-
cave. The shape of the processus orbitalis is similar to that
of extant Todidae, and this process is proportionally shorter
than in crown group Momotidae. As in the latter but con-
trary to modern todies, it bears a marked flange near its
dorsal margin (Fig. 4b; crista orbitalis of Elzanowski et al.
2000).
Fifteen praesacral vertebrae are preserved, as well as
four caudal vertebrae and the pygostyle. The atlas seems to
be lost. The processus spinosus and the zygapophyses
caudales of the axis are more prominent than in extant
Todidae. The third cervical vertebra is seen in cranial view,
but is poorly preserved; the other cervical and thoracic
vertebrae likewise do not exhibit features of taxonomic or
phylogenetic significance. The lamina pygostyli is some-
what wider than in extant Todidae.
Coracoid morphology of Palaeotodus is well known
(Mayr and Knopf 2007b), but the scapula is for the first
time completely preserved in a Palaeotodus specimen. It
differs from the scapula of crown group Todidae in that
the corpus is less angled (Fig. 4g, h), which by com-
parison with the Momotidae is probably the plesiomor-
phic condition. The sternum shows the dorsal surface
(Fig. 4l); the trabeculae of the right side and the caudal
portion of the left trabecula lateralis are broken. As in the
previously reported specimen of the Frauenweiler tody
(Mayr and Knopf 2007b), three processus costales can be
counted.
Fig. 3 Palaeotodus itardiensis Mourer-Chauvire, 1985 from the
early Oligocene of Frauenweiler. Specimen HLMD-WT 590 (coated
with ammonium chloride) with interpretative drawing. Abbreviations:
axs axis, cdv caudal vertebra, fsv fish vertebrae, hal hallux, lcm left
carpometacarpus, lco left coracoid, lfe left femur, lhu left humerus,
lqu left quadrate, lra left radius, lsc left scapula, ltb left tibiotarsus,
ltm left tarsometatarsus, lul left ulna, mdb mandibular ramus, pelpelvis, phl pedal phalanges, pyg pygostyle, r rib, rco right coracoid,
rfe right femur, rhu right humerus, rqu right quadrate, rsc right
scapula, rtm right tarsometatarsus, rul right ulna, ste sternum, vvertebra
New specimens of Eurotrochilus and Palaeotodus 391
123
The left humerus is visible in cranial view, whereas the
right exposes the caudal surface, which cannot be seen in
the skeleton described by Mayr and Knopf (2007b). As
in crown group Todidae, the caput humeri is caudally
inflected. The proximal end of the ulna of the Frauenweiler
tody is, for the first time, well preserved in the new speci-
men (Fig. 4k). With a width of 2.8 mm, it has the same size
as the holotype ulna of P. itardiensis (Mourer-Chauvire
Fig. 4 Selected postcranial bones of Palaeotodus itardiensis Mourer-
Chauvire, 1985 from the early Oligocene of Frauenweiler (HLMD-
WT 590). a, b left (a) and right (b) quadrate in lateral view, in
comparison to c the right quadrate of the Blue-crowned Motmot,
Momotus momota (Momotidae); d axis in cranial view; e pygostyle in
lateral view; f left coracoid in dorsal view; g right scapula in medialview, in comparison to h the right scapula of the Cuban Tody, Todusmulticolor (Todidae); i right humerus, right coracoid, distal end of left
tibiotarsus, and proximal right tarsometatarsus; j left humerus in
cranial view; k proximal end of left ulna in cranial view; l sternum in
dorsal view; m right femur in caudal view; n pelvis in ventral view.
Fossil bones are coated with ammonium chloride. Abbreviations: cdmcondylus medialis, cla condylus lateralis, cph caput humeri, cro crista
orbitalis, cst constriction, ctd cotyla dorsalis (broken), ctv cotyla
ventralis, cvt concavity distal of tuberculum ligamenti collateralis
ventralis, ltb left tibiotarsus, olc olecranon, pit pit at insertion area of
musculus biceps brachii, pot processus oticus, psp processus spinosus,
rco right coracoid, rdg ridge, rhu right humerus, rtm right
tarsometatarsus, spe spina externa, tcv tuberculum ligamenti collat-
eralis ventralis, tim trabecula intermedia, tla trabecula lateralis, zyczygapophysis caudalis. Scale bars equal 2 mm for a–e, and 5 mm
for f–n
392 G. Mayr, N. Micklich
123
1985), which it also closely resembles in morphology
(compare Fig. 4k with Mourer-Chauvire 1985: pl. 1). As in
P. itardiensis and extant Todidae, the cranial surface of the
shaft, just distal of the tuberculum ligamenti collateralis
ventralis, is deeply concave. The cotyla dorsalis is broken,
but the shape of the cotyla ventralis and the well-delimited
olecranon match the holotype of P. itardiensis. As in the
latter, there is a marked oblique muscle attachment scar
distal of the cotyla ventralis (Fig. 4k). The tuberculum
supracondylare ventrale is somewhat larger than in crown
group Todidae and Momotidae. The left carpometacarpus
shows the previously unknown ventral surface. The pro-
cessus extensorius is larger than in crown group Todidae,
but otherwise the bone is too poorly preserved for the rec-
ognition of osteological details.
The pelvis is visible in ventral view and lacks the left
ischium and ilium (Fig. 4n). As in extant Todidae, ten
vertebrae are fused in the synsacrum. The left femur is
exposed in caudal view (Fig. 4m) and has similar propor-
tion to the femora of Todidae and Momotidae, whereas this
bone is stouter in Alcedinidae and Meropidae. As in
Momotidae and most other birds, the crista tibiofibularis of
the trochlea fibularis reaches farther proximally than the
crista supracondylaris lateralis, whereas these cristae have
the same proximal extent in crown group Todidae. The
distal end of the left tibiotarsus is visible in caudal view
(Fig. 4i). The trochlea cartilaginis tibialis is somewhat
narrower and proximo-distally deeper than in crown group
Todidae. Of the tarsometatarsi, only the proximal half
(right) and proximal tip (left) are preserved. The surface of
the right tarsometatarsus is unusually eroded and porous
and does not allow the recognition of any details (Fig. 4i);
because other bones, i.e. the distal end of the tibiotarsus
and the proximal left tarsometatarsus, do not indicate a
juvenile condition of the specimen, we consider this mor-
phology to be either a taphonomic artifact or of pathologic
origin. A few pedal phalanges are scattered over the slab.
Among these is the proximal end of the first phalanx of the
right hallux, which bears the lateral projection previously
identified as an alcediniform apomorphy (Mayr 1998; Mayr
and Knopf 2007b).
Discussion
The composition of the Frauenweiler avifauna appears
quite unbalanced, because all specimens of non-marine
birds belong to very small species. The only larger avian
fossils are from the procellariiform taxon Diomedeoides
and the loon Colymboides (Mayr et al. 2002; Mayr 2004a,
2009a), whereas remains of medium-sized or large terres-
trial birds have not been found so far. Diomedeoides is also
the most abundant avian taxon, which is not surprising
because the Frauenweiler sediments were deposited in a
marine environment. It is more unusual, however, that the
second most abundant avian species is the stem group
hummingbird Eurotrochilus inexpectatus. The chance that
fossils are overlooked is much higher for such tiny species,
and absence of larger birds in the Frauenweiler site is
certainly a taphonomic and not a collecting artifact. Pos-
sible, albeit still to be verified, explanations for the lack of
larger terrestrial birds may be differences in the drifting
distances of differently sized bird carcasses, or a higher
susceptibility of smaller birds to be blown into the open sea
by storms.
The preservation of the new specimens reported here
and that of other small birds from Frauenweiler (e.g., Mayr
2000, 2004a, b, 2005b, 2006; Mayr and Manegold 2006) is
remarkable, because the fossils are completely disarticu-
lated, with the bones being jumbled but nevertheless clo-
sely associated. This curious preservation contrasts with
that of fish skeletons from the fossil site, which are often
fully articulated (Micklich et al. 2009). Clearly, disarticu-
lation of the bird skeletons must have occurred after the
carcasses sank to the seafloor, but before they were
embedded in the sediment. Subsequent disturbance of the
bones may have been due to scavengers such as small
crustaceans, but the conditions on the ground of the Rup-
elian sea at the Frauenweiler locality are considered dys- or
poikiloaerobic, i.e., hostile for benthic macroorganisms
(Grimm et al. 2002; Micklich and Hildebrandt 2005;
Micklich et al. 2009). Alternatively, thus, the carcasses
may have exploded due to the formation of putrefaction
gases, or the bones could have been mingled by fluctuating
ground currents. The latter, however, cannot have been
very strong, as the fish skeletons do not show special pat-
terns of alignments. The preservation of the new Eurotro-
chilus specimen, whose bones are closely clustered in a
well-defined area, is suggestive of a pellet, either from an
avian predator or a fish that ingested the floating or sinking
bird and regurgitated the indigestible parts. For a well-
founded explanation of the taphonomy of small birds from
Frauenweiler, further actuopalaeontological studies of the
decay of bird carcasses in marine environments are
desirable.
Nannoplankton stratigraphy places the Frauenweiler
‘‘Fischschiefer’’, from which most bird fossils stem, into
the unit NP 23 (Grimm et al. 2002). However, because
very few mammalian remains are known from the locality,
and these have not yet been studied in detail, exact bio-
stratigraphic correlation of the Frauenweiler sediments
with other terrestrial fossil sites has proven difficult
(in addition to two bats, a mandible of a creodont was
recently found; Monninger and Frey 2009). The Quercy
specimens of P. itardiensis stem from the site Itardies,
whose sediments belong to the Mammalian Paleogene unit
New specimens of Eurotrochilus and Palaeotodus 393
123
MP 23 (Mourer-Chauvire 1995), and occurrence of this
species in Frauenweiler suggests an absolute age of about
31 million years for the fossiliferous sediments of the
locality (Legendre and Leveque 1997).
The Frauenweiler fossil site is certainly among the most
important European localities for Rupelian bird fossils.
Unfortunately, the future of the locality is insecure, and
most areas of the former pit have been used as a landfill
during the past years. There are ongoing negotiations with
the current owners to ensure further excavations, and we
are confident that future bird fossils will continue to con-
tribute to a better understanding of a poorly known period
of the Cenozoic evolution of birds.
Acknowledgments We thank Harald and Annette Oechsler for
donating the Eurotrochilus specimen to SMNK, and Klaus Weiß for
the excellent preparation of both fossils—without their tireless
enthusiasm many of the Frauenweiler fossils would not have been
unearthed. We further thank Dino Frey for the loan of the Eurotro-chilus specimen, and Sven Trankner for taking the photographs.
Ludwig Hildebrandt made the new profilings, which established the
base for a precise stratigraphic allocation of the fossils. Recent
excavations in the Frauenweiler fossil site were funded by the
National Geographic Society (8553-08). The comments of two
anonymous reviewers improved the manuscript.
References
Baumel, J.J., and L.M. Witmer. 1993. Osteologia. In Handbook ofavian anatomy: Nomina Anatomica Avium, eds. J.J. Baumel,
A.S. King, J.E. Breazile, H.E. Evans, and J.C. Vanden Berge.
Publications of the Nuttall Ornithological Club 23: 45–132.
Bochenski, Z., and Z.M. Bochenski. 2008. An Old World humming-
bird from the Oligocene: A new fossil from Polish Carpathians.
Journal of Ornithology 149: 211–216.
Elzanowski, A., G.S. Paul, and T.A. Stidham. 2000. An avian
quadrate from the late Cretaceous lance formation of Wyoming.
Journal of Vertebrate Paleontology 20: 712–719.
Grimm, K.I., M.C. Grimm, A. Kothe, and T. Schindler. 2002. Der
,,Rupelton’’ (Rupelium, Oligozan) der Tongrube Bott-Eder bei
Rauenberg (Oberrheingraben, Deutschland). Courier Fors-chungsinstitut Senckenberg 237: 229–253.
Karhu, A. 1988. Novoye semeystvo strizheobraznykh iz paleogena
Yevropy [A new family of swift-like birds from the Paleogene of
Europe]. Paleontologicheskii Zhurnal 3: 78–88.
Karhu, A. 1999. A new genus and species of the family Jungorni-
thidae (Apodiformes) from the late Eocene of the Northern
Caucasus, with comments on the ancestry of hummingbirds. In
Avian paleontology at the close of the 20th century: Proceedingsof the 4th International Meeting of the Society of AvianPaleontology and Evolution, Washington, D.C., 4–7 June1996, ed. S.L. Olson. Smithsonian Contributions to Paleobiology89: 207–216.
Legendre, S., and F. Leveque. 1997. Etalonnage de l’echelle
biochronologique mammalienne du Paleogene d’Europe occi-
dentale: Vers une integration a l’echelle globale. In Actes duCongres BiochroM’97, eds. J.-P. Aguilar, S. Legendre, and J.
Michaux. Memoires et Travaux de l’Ecole Pratique des HautesEtudes, Institut de Montpellier 21: 461–473.
Louchart, A., N. Tourment, J. Carrier, T. Roux, and C. Mourer-
Chauvire. 2008. Hummingbird with modern feathering: an
exceptionally well-preserved Oligocene fossil from southern
France. Naturwissenschaften 95: 171–175.
Mayr, G. 1998. ‘‘Coraciiforme’’ und ‘‘piciforme’’ Kleinvogel aus dem
Mittel-Eozan der Grube Messel (Hessen, Deutschland). CourierForschungsinstitut Senckenberg 205: 1–101.
Mayr, G. 2000. A new mousebird (Coliiformes: Coliidae) from the
Oligocene of Germany. Journal fur Ornithologie 141: 85–92.
Mayr, G. 2004a. A partial skeleton of a new fossil loon (Aves,
Gaviiformes) from the early Oligocene of Germany with
preserved stomach content. Journal of Ornithology 145: 281–286.
Mayr, G. 2004b. Old World fossil record of modern-type humming-
birds. Science 304: 861–864.Mayr, G. 2005a. New trogons from the early tertiary of Germany. Ibis
147: 512–518.
Mayr, G. 2005b. A tiny barbet-like bird from the lower Oligocene of
Germany: The smallest species and earliest substantial fossil
record of the Pici (woodpeckers and allies). The Auk 122: 1055–
1063.
Mayr, G. 2006. First fossil skull of a Paleogene representative of the
Pici (woodpeckers and allies) and its evolutionary implications.
Ibis 148: 824–827.
Mayr, G. 2007. New specimens of the early Oligocene Old World
hummingbird Eurotrochilus inexpectatus. Journal of Ornithol-ogy 148: 105–111.
Mayr, G. 2009a. Notes on the osteology and phylogenetic affinities of
the Oligocene Diomedeoididae (Aves, Procellariiformes). FossilRecord 12: 133–140.
Mayr, G. 2009b. Paleogene fossil birds. Berlin: Springer.
Mayr, G., and C. Knopf. 2007a. A stem lineage representative of
buttonquails from the lower Oligocene of Germany—fossil
evidence for a charadriiform origin of the Turnicidae. Ibis 149:
774–782.
Mayr, G., and C. Knopf. 2007b. A tody (Alcediniformes, Todidae)
from the lower Oligocene of Germany. The Auk 124: 1294–1304.
Mayr, G., and A. Manegold. 2004. The oldest European fossil
songbird from the early Oligocene of Germany. Naturwissens-chaften 91: 173–177.
Mayr, G., and A. Manegold. 2006. New specimens of the earliest
European passeriform bird. Acta Palaeontologica Polonica 51:
315–323.
Mayr, G., D.S. Peters, and S. Rietschel. 2002. Petrel-like birds with a
peculiar foot morphology from the Oligocene of Germany and
Belgium (Aves: Procellariiformes). Journal of Vertebrate Pale-ontology 22: 667–676.
Micklich, N. 1998. New information on the fish fauna of the
Frauenweiler fossil site. Italian Journal of Zoology 65(supple-
ment): 169–184.
Micklich, N., and L. Hildebrandt. 2005. The Frauenweiler clay pit
(‘‘Grube Unterfeld’’). Kaupia: Darmstadter Beitrage zur Na-turkunde 14: 113–118.
Micklich, N., and N. Parin. 1996. The fishfauna of Frauenweiler
(Middle Oligocene, Rupelian; Germany): First results of a
review. Instituto Espanol de Oceanografıa, PublicacionesEspeciales 21: 129–148.
Micklich, N.R., J.C. Tyler, G.D. Johnson, E. Swidnicka, and A.F.
Bannikov. 2009. First fossil records of the tholichthys larval
stage of butterfly fishes (Perciformes, Chaetodontidae), from the
Oligocene of Europe. Palaontologische Zeitschrift 83: 479–497.
Monninger, S., and E. Frey. 2009. Oligocene bats from the clay pits
around Frauenweiler. In 7th annual EAVP meeting 2009,abstract volume, eds. D. Schwarz-Wings, O. Wings, and F.
Sattler. Aachen: Shaker.
Mourer-Chauvire, C. 1985. Les Todidae (Aves, Coraciiformes)
des Phosphorites du Quercy (France). Proceedings of theKoninklijke Nederlandse Akademie van Wetenschappen, SeriesB 88: 407–414.
394 G. Mayr, N. Micklich
123
Mourer-Chauvire, C. 1995. Dynamics of the avifauna during the
Paleogene and the early Neogene of France. Settling of therecent fauna. Acta zoologica cracoviensia 38: 325–342.
Olson, S.L. 1976. Oligocene fossils bearing on the origins of the
Todidae and the Momotidae (Aves: Coraciiformes). SmithsonianContributions to Paleobiology 27: 111–119.
New specimens of Eurotrochilus and Palaeotodus 395
123