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.

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