Of the Clay Pit Unterfeld at Rauenberg (Rhein-Neckar-Kreis, Baden-W

Home , Apterodon, Creodonta

110114_kaupia17.05.qxd.05.vrz 16.01.201118:05UhrSeite107 sich morphologisch vonsich morphologisch denbeidenbisherausEuropa bekannten Formen von nachgewiesen: folge) deraufgelassenenTongrube Unterfeld be e udtle möglichweise imBereich einerFlussmündung. der Fundstelle, in dieMP-Zonen VorkommenDas derGattung close to ariver mouth. probably thesedimentsatfindingplace, theshoreline of The specimenproves of theimmediate vicinity Badré &Böhme, Landsäugetie Mit demFund einesrechten Unterkieferfragmentes mitTeilen derBezahnung wurde zumersten Mal ein Zusammenfassung Erbprinzenstrasse Abteilung, Adressen der Autoren: Baden-Württemberg): at (Rhein-Neckar-Kreis, of theClayRauenberg PitUnterfeld First creodont carnivore from theRupelianClays (Oligocene) Michael Morlo &Mandana Hensel Madelaine Böhme, Wolfgang Munk, Frey, Eberhard morphologically differs frommorphologically Germany): Southern berg, theopencastClay PitUnterfe Graue Schichtenfolge) of (Rupelian, theFishShalesubformation Mammalia terrestrial depositsof inthemarine for thepresence of dentitionforthefirsttimeproves evidence includingafragmentary mandibularramus aright The discovery of Abstract & Böhme, emn,[email protected]; Germany, Erbprinzenstrasse Geowissenschaftliche Abteilung, Museum fürNaturkunde Karlsruhe, D-60261 [email protected]; D-72076 width. The occurrence of Theoccurrence width. emn,[email protected]. Germany, rnfr,Gray [email protected]; Germany, Frankfurt, Tübingen andSenckenberg Center for 2005 r in den marinen r indenmarinen Ablagerungen derFischschiefer-Subformation (Rupel Apterodon rauenbergensis .gaudryi and A. 2005 22 bis 23 .gaudryi and A. Eberhard FreyEberhard Apterodon u DerFund belegtdieunmittelbare Ufernähe des Ablagerungsraumes imBereich zu. Apterodon rauenbergensis the othertwo of species 1 , University Tübingen, Geological Insitute, Sigwartstrasse Sigwartstrasse Insitute, Geological University Tübingen, Madelaine Böhme, ,D- Apterodon Fischer, 76133 Michael Morlo Michael losfra dnicto fthelayers ascoming from allows foranidentificationof Fischer, talce uemfrNtrud alrh,Geowissenschaftliche Staatliches Museum fürNaturkunde Karlsruhe, , Apterodon rauenbergensisApterodon alrh,Gray [email protected]; Germany, Karlsruhe, 1880 .s.(root,Handnia) DasFundstück unterscheidet Hyaenodontidae). (Creodonta, sp. n. lässt eine säugetierstratigraphische Eingrenzung derFundschichtenlässt einesäugetierstratigraphische ai,dassp4 darin, 1880 i Rauenberg (Rhein-Neckar-Kreis, Baden-Württemberg) Baden-Württemberg) i Rauenberg(Rhein-Neckar-Kreis, Human Evolution andPalaeoecology ( oshnsntttSnkneg Senckenberganlage Forschungsinstitut Senckenberg, , in having p4,m .p Cedna yeootde.Thespecimen Hyaenodontidae). (Creodonta, n.sp. Apterodon da aebr RenNca-ri,Baden Württem- ld atRauenberg(Rhein-Neckar-Kreis, Mandana Hensel,Weinweg ,m 1 und m known fromknown Europe 1 and m 2 etwa gleich langundbreitetwa gleich sind. 2 fnearlythesamelengthand of .intermedius A. Staatliches Wolfgang Munk, 1 ,D- ium, Graue Schichten- Graue ium, n.sp. hep), .intermedius A. 1 ,D- 76327 76133 Lange-Badré mp Pfinztal, 22 Karlsruhe, or Lange- mp 25 , 10, 23.

Figure 1: Geography and strati- graphy of the clay pit Unterfeld; A: Location of the clay pit, B: ﬁnding place within the clay pit, C: schematised stratigraphical column of the German Oligocene, the asterisk marks the ﬁnding position of SMNK PAL 6524.

C B

500 m

During the emergency excavation in the year 2007 the Introduction fragment of a right mandibular ramus was discovered Since Roman times the Rupelian clays (Early in the debris along a dirt path in the north eastern Oligocene) in the northern Baden area (Baden-Würt- corner of the clay pit Unterfeld (Fig. 1 A) leading to temberg) were exploited for the production of bricks the excavation area by the private collector Klaus- and concrete. It was in the early seventies when the Dieter Weiß (Fischbach, Hesse State). He pre-prepared sediment became recognised for their fossil content. the specimen and handed it over to smnk.After a Until today more then 70 taxa of teleost fishes and final cleaning, the specimen was incorporated into the chondrichthyans have been described with Aeoliscus palaeontology collection of the smnk under the and Sardinites being the most abundant genera number smnk pal 6524. It represents the first creodont (Micklich 1998, Micklich & Parin 1996, Micklich & record from the area and thus the first evidence for a Hildebrandt 2005). In some layers, the sediment is terrestrial tetrapod in the Northern Badish Rupelium. full of plant debris, cuticula preservation and foraminifera. Seaweed, gastropods, bivalves, crusta- It should be mentioned here that more that 80% of tions and fossil logs are comparably rare (Grimm et all finds registered in public collections come from al. 2002), as are sea turtles (Alexander & Frey, this private collectors. Without them the fossil treasures volume), which are only known from a handful of of the Northern Badish Rupelian clay pits could never specimens. Plants are not the only remnants from the have been secured to such an amount for the scien- life of the coastal back lands. A few species of birds tific community. As representatives for all the those have been described, whereby typical coastal or sea- enthusiasts that worked in the clay pits over years we birds are rarer than typical inland birds (Mayr et al. like to dedicate this article to Klaus-Dieter Weiß as 2002, Mayr 2004b). The fossil record of the North well as to Annette and Harald Oechsler (Paläo-Geo e.V.) Baden Rupelian yielded trogons as well as Europe’s for their unselfish contributions to science. oldest songbirds and woodpeckers (Mayr 2005). The bird fossil with the highest impact factor form the clay pit Unterfeld near Rauenberg is a fossil Material and methods humming bird, Eurotrochilus inexpectatus, which The specimen smnk pal 6524 was mechanically pre- proved an early origin of the group in the Old World pared and photographed with an Olympus E620 (Mayr 2004 a, 2007). The insect assemblage is (wide angle zoom 14–42 mm) in macro mode. The dominated by Coleoptera and Hemiptera. Other taxa, image was composed with CorelDraw 12 and X3.The like Hymenoptera, Amphiesmenoptera (Lepidoptera schematic drawings were created with Corel Trace. + Trichoptera), Diptera and Mantodea are only known from single specimens (Monninger & Frey, this volume). Remnants of mammals are generally rare as well. From what is guarded in public collections there are a few fragments and one single sub-com- plete skeleton of Sirenia (cf. Halitherium) and two Microchiroptera, which both are under description. Today, the last open clay pit with palaeontological excavation activites is the Clay Pit Unterfeld, territory of Rauenberg (Fig. 1A). 110114_kaupia17.05.qxd.05.vrz 16.01.201118:05UhrSeite109 foraminifera ( foraminifera brandt this hasnotyet published. beenformally but have to to beraised order level, and Oxyaenidae, Hyaenodontidae thetwo includedfamilies, quence, annual averageclima with temperatures between Thisrefers to aCfa subtropical climate. a humid, Europeanthe northern Oligocene wasdominated by re:›Creodonta‹ Order: According to Superorder: Laurasiatheria Laurasiatheria Superorder: depth of authorssuggestawater Some cussed controversially. thesedimentsisstilldis- The depositiondepthof distances.in irregular occur carbonates silty bandsof grey light thin, stone, Within thismud- water marlymudstone. rich ered, predominantly well lay- dark-grey, mostly consists of FishShalesub-formation The tocene covering layers. theFishShalesbelow thePleis- of the bottom part ls:Mammalia Class: Systematik palaeontology ( and 18 Grimm Trunko &Munk according sectionof to thegeological likelyvery derives from alayer notbeyond layer thespecimen which wasrecorded by thefinder, tion, From thefindingposi- theclay pitUnterfeld. der of shoul- weathered thestrongly north-eastern of debris wasdiscoveredThe mandibularfragment inthe andpalaeoenvironmentGeology ( Hoppe be notvalid(e.g. diphyletic by authorsandthus believed allmodern to theorder ›Creodonta‹ isregarded as Todate, Remark: nrcas Eutheria Infraclass: ,while 2002), middle-sized predators like anidealhabitatforsmallto byly interrupted rivers, like- palmsandrarelyconifers, laurels, dense forest of thebacklandwascovered a with litter, andleaf debris brücken dip( Rauenbergsouthboundinto theLangen- of margin Wiesloch andcontinues alongthewestern of the city the Palaeogene today of passestheeastern margin Thiscoastline faultthatalready existed in main fault. the toby parallel theeastof another faultrunning wasformed Thecoastline itself about onekilometre. shallow of water area awidth distalto thisfaultwith likely to belongs a clay The pititself the findingplace. approximately runs escarpment 250 the Themainfaultof juveniles. and theabundancyof many shallow water fishtaxa on theoccurrence of ftheRupelian includingtheentire of overburden theprevious theclay pit, western area of In contrast to the the FishShalesub-formation. eroded away ( era thepre-glacial during theFish Shaleswas mation andthetop ten meters of Köppen Fig. 2002, n naeaerifl of andanaverage rainfall ° 1993). ta.(2002) shows theexclusive presence of et al. ( 100 2005) suggestashallow water based regime .A scnlddb nso os plant logs, As isconcluded by findsof 1931). 1 ) In any casethespecimencomes from B). Weiler to to Pross Thürach Trunko &Munk 200 ). The geological sectionof Thegeological 1998). ei Morlo & Lewis Linné, Huxley, metres basedonbenthontic Cope, ) the middle part of (1998)themiddlepart et al. ( 1966) and 1904, 1758 1875 Murphy, Apterodon. 1889 Doebl 1998, Micklich &Hilde- 1000 .As aconse- 2010). 1976 metres eastof Meletta Grimm Grimm 2001 Grimm mm to Doebl , Foellmer & sub-for- et al. 1300 et al. 1976 11 16 mm , ° 1 ( specimenisidentifiedas The : Diagnosis ous otherauthorsis»Frauenweiler«. aiy Hyaenodontidae Family: Genus: ufmle Apterodontinae Subfamilie: agno p of margin mandibularsymphysis thecaudal level with of ( (Fischer ( (as intheother species African butinthisspeciesp 1930), Holotypus Rauenberg. of theCity of Clay The PitUnterfeld liesontheterritory the future. the scientificexcavations intheClay PitUnterfeld for which wasmuch engagedto secure government of the city Rauenberg, of theCity inhonourof berg; Derivatio nominis Derivatio Species: mer species(Tab. medius tive to m a newspecies: pal size thenewspecimensmnk we attribute than m (Fig. was aboutfourtimesaslongitsmaximum height ing to ( p name aspublishedby Theofficiallocality SouthGermany. North Baden, Clay PitUnterfeld atRauenberg, Early Oligocene, Locus typicus für Naturkunde Karlsruhe. theStaatliches Museum collectionspalaeontology of p asmassivetwice asparaconid asreconstructed from e etso millimetres (Fig. a fewtenths of thevaluemay differ even if same lengthandwidth, molar length (visible inp molar length(visible talonidonethird of the crowns isnotpreserved, eaielnt fm relative lengthof mkpal smnk ( includingtheEuropean taxa species, Fig. , Andrews Fischer ios&Gingerich & Simons Böhme & Lange-Badré 1964), 4 3 and m molariform (Fig. molariform 2 3 Andreae A, A’, B, B’). Further diagnostic features diagnostic Further accord- B’). A’, B, A, .Dueto thesedifferences inrelative tooth ). 1 Lange-Badré &Böhme Apterodon the specimenappearsto becloserto thefor- , and the same is true for andthesameistrue , peoo rauenbergensis Apterodon 2 , , 1 eas fissotmniua which mandibula, itsshort 1830) becauseof inp 1880) 6524 .intermedius In A. ). saghensis andA. 1905) and onlyspecimen: .gaudryi in A. (Fig. 3 ihSaesbfrain Rupelian, FishShalesub-formation, : (Fig. .rauenbergensis. A. differs from allother 2 ( 1887), A, A’, C, C’), however, the height of theheight however, C’), A’, C, A, : 1 Fischer 2 Fig. , rauenbergensis 1 4 ,B) p B’); B, /m 2 ,m , Grimm ,A,C ’;protoconid almost C’); A’, C, A, 1976 2 Matthew , 3 in comparison to 1 A. altidens ( altidens is A. ). A. gaudry gaudry 2005)and A. Leidy, 3 and m 4 , ) them and m 1880 is clearlylongerthanm Szalay 3 et al. with cingulid; with mkpal smnk ( .macrognathus A. 1869 r:caudalend 2005) are: 2 2 As p As 1 .gaudryi A. ( having aboutthe 2 lt)=o Rauen- (lat.) =of ,C) Closestin C’). C, Fig. ; .sp. n. ,Dashzeveg 1909), 2002 , is clearlylonger 1967 Apterodon .intermedius A. 4 Schlosser is largerrela- 2 Apterodon and numer- 6524 ,A,C C’). A’, C, A, .inter- A. (see Tab. , 6524 , to 1

109 E. Frey,W. Munk, M. Böhme, M. Morlo & M. Hensel | First creodont carnivore from the Rupelian Clays of the former claypit Unterfeld at Rauenberg: Apterodon rauenbergensis n.sp. | 110114_kaupia17.05.qxd.05.vrz 16.01.2011 18:05 Uhr Seite 110

Figure 2. Apterodon rauenber- cingulum gensis n.sp. SMNK PAL 6524: foss. mass. m2 m1 p4 p3 p2 p1 A, A’: lateral view, proc. ang. B, B’: Medial view, C, C’: occlusal view. foss. mass. = fossa masseterica, for. ment. = foramen mentale, proc. ang. = processus angularis, symph. = symphysis, m = molar, p = premolar. A A’ for. ment.

p2 p3 p4 m1 m2

symph.

B B’ proc.ang.

proc.ang. m2 m1 p4 p3 p2 p1

C C’ for. ment.

30 mm fracture surface enamel root

Description the specimen and bears a protoconid which is about The fragment of a right mandibular ramus has a total twice the size of the paraconid in occlusal view, length of 88 mm (Fig. 2). The processus coronoideus whereas the paraconid is mostly eroded. The tooth is including the caput mandibulae and the processus 8 mm long, 5 mm high and at the crown base has a angularis are broken off along an oblique line, running width of 4.5 mm. P3 lacks the entire crown but both from the cranial base of the processus coronoideus roots are in situ showing circular cross-sections. The at an angle of about 45° caudoventrally through the tooth had a length of about 10 mm as is estimated fossa masseterica to the ventral margin of the dentary from the space between p2 and p4.P4 also preserves and terminates at the base of the processus angularis two roots with circular cross-sections. The crown is (Fig. 2 A, A’). The ventral half of fossa masseterica extremely eroded but apparently has been tricuspid. is preserved with a triangular piece of bone missing at The protoconid likely was the biggest of the three the caudodorsal fracture line (Fig. 2 A, A’). The lateral cusps. Labially, the trace of a cingulid is visible. The wall of the base of the processus angularis is missing. tooth has a length of 12.3 mm and a maximum width Rostrally, the madibular ramus has broken at the at the mid crown height is 6.6 mm. The double rooted caudal terminus of the mandibular symphysis with a m1 lacks the entire crown. Both roots are circular fracture line running caudoventrally at an angle of in cross-section. The tooth fragment is 12.0 mm long about 50° to the dental margin of the mandibular at the break along the roots. M2 is preserved similar ramus (Fig. 2). Of the mandibular symphysis the cau- to m1 with most of the crown missing. The length dal-most part is visible, terminating ventral to the measured along the breakage face of the roots, which caudal margin of p3 (Fig. 2 B, B’). The highest dorso- are circular in cross section, is about 12.4 mm. At ventral expansion of the specimen with 31 mm lies the base of the crown both teeth were certainly a little level with the fracture caudal to m2. On the labial longer. M3 is completely missing and not even a face the caudal foramen mentale opens ventral to the trace of the rostral border of its alveolus is visible. This caudal root of p3 at about the mid height of the suggests that the specimen comes from a juvenile mandibular ramus (Fig. 2 A, A’). The caudal margin individual with all permanent premolars and molars of a second foramen mentale is visible 20 mm further erupted except m3. However, the eruption sequence rostrally and is cut half by the rostral fracture. It lies of apterodontines remains unknown, even though it ventral to the middle of p2 and level with the dorsal is aparent that m3 and the canine are the latest erupt- third of the mandibular ramus. ing teeth in hyaendontines (Mellett 1977).

Incisivi and canini are missing due to the loss of the symphyseal part of the mandibular ramus. All other teeth are preserved with their crowns strongly abrad- ed (Fig. 2 A, A’,C, C’). Of p2 only an indistinctive fragment of the root is preserved. p3 has two roots. The crown is the best preserved of the dental series of 110114_kaupia17.05.qxd.05.vrz 16.01.2011 18:05 Uhr Seite 111

Figure 3: Plots of selected tooth ratios in Apterodon. A) ratio m1/m2, B) ratio p4/m1.

A B

◆ A. altidens A. gaudryi A. intermedius ✕ A. rauenbergensis n. sp. ▲ A. saghensis A. macrognathus

Taphonomy and diagenesis Discussion The specimen was embedded within a concretion- According to Lange-Badré & Böhme (2005) the genus like mudstone nodule. Only the ventral ridge of the Apterodon consists of five valid species, two of which 1880 mandibular ramus was visible, when it was collected. come from Europe: A. gaudryi Fischer, Rauenberg: n.sp. rauenbergensis Apterodon | Therefore it represents a primary fragment. The abra- (syn. A. flonheimensis Andreae, 1887; Lange-Badré & sion of the teeth and the margins of the fractures sug- Böhme 2005), A. intermedius Lange-Badré & Böhme, gest, that the specimen has been transported to the 2005, and three are reported from Egypt: A. macrog- place of deposition, either by a near-by river, where nathus Andrews, 1905, A. altidens Schlosser, 1910, the animal died a few kilometres upstream and trav- A. saghensis Simons & Gingerich, 1976. elled downstream already isolated. Another possibili- Because of the fragmentary fossil record the taxono- ty would be that the animal died at the shore and was my of the genus remained controversial for a long dismembered by the action of the surge. In both cases time (for a review see Langer-Badré & Böhme 2005). scavengers may have initially dismembered the car- Until today the fragmentary preservation states of cass. Because the massive bone would not have drift- Apterodon remnants in Europe make a diagnosis to ed at the water surface, it must have been transported species level problematic, because many characters to the final embedding place by currents. A final short cannot be directly compared with each other on distance transport from the shore by a seabird (e.g. species level. Until further finds, the diagnostic differ- Diomedoides brodkorbi Mayr, Peters & Rietschel ences are extremely critical with regard to statistical 2002) would also be an option to explain the arrival evaluations but still are justified on the level of dis- of the specimen at the embedding place. tinct morphotypes (see e.g. Lange-Badré & Böhme 2005). The specimen presented here differs from the During late diagenesis some compaction breaks were hitherto described European and African species in either caused by sediment compaction or possibly the ratios of functionally crucial teeth: p4 and the two tectonical compression or both. This compaction molars (see Tab. 1, Fig. 3). Give the functional impe- caused a deep shear fracture that runs parallel to the tus of the masticatory apparatus of apterodontines ventral margin of the mandibular ramus. The speci- we feel it justified to erect a new morphotype that men finally was impregnated with iron sulfide and formally adds a third Apterodon species to the Euro- very likely is with pyrite. pean inventory of this genus. However, further dis- coveries may yield more data that may show the plas- ticity of the masticatory apparatus of Apterodon. Such data are only available for A. macrognathus which shows a variation of p4/m1-length ratio between 1.27 and 1.40 (n = 4 specimens; Fig. 3 B) and of m1/m2-length ratio between 0.72 and 0.79 (n = 4 specimens, Fig. 3 A). The European branch of the genus Apterodon is restricted to mp 22 and 23, which means that for the first time the Rupelian Clays in the Rauenberg-Wiesloch area are referable to a comparably narrow mammalian biozone. E. Frey,W. Munk, M. Böhme, M. Morlo & M. Unterfeld claypit at of the former from the Rupelian Clays creodont carnivore Hensel | First 111 110114_kaupia17.05.qxd.05.vrz 16.01.2011 18:05 Uhr Seite 112

Figure 4: Apterodon rauenbergensis n.sp., temptative reconstruction as a short-faced, semi-aquatic predator.

Table 1: Comparative mesurements of ﬁve species of Apterodon according to LANGE-BADRÉ & BÖHME (2005).

So far, the European fossils of Apterodon are coming have been interpreted as bone/meat feeder (Morlo, from three sites: the fissure fillings of the Phospho- Gunnell & Nagel 2010). The habitat of the short- rites du Quercy (Dept. Lot, France; a juvenile individ- faced Apterodon may thus may have been the imme- ual of A. gaudryi, Fischer, 1880), and the coastal- diate vicinity of large rivers or even coastal-marine marine sands of Flonheim (Mainz Basin; A. gaudryi, settings. syn.: A. flonheimensis, Andrae, 1987) and Espenhain (Weisselster Basin; A. intermedius, Böhme, 2001). The new find from Unterfeld again originate from coastal- Acknowledgements marine sediments, which opens the question about Herr Klaus-Dieter Weiß (Fischbach/Ts.), mit dem be- the possibility of a semi-aquatic (otter-like, see also reits seit vielen Jahren eine äußerst fruchtbare the temptative reconstruction in Fig. 4) lifestyle of Zusammenarbeit besteht, machte den Fund im Sep- Apterodon. However, no characters typical for a semi- tember 2007 und übereignete ihn danach dem smnk. aquatic lifestyle are known from postcranial remains Das Fundstück wurde von Herrn René Kastner of Apterodon (MM, unpubl. data). The peculiar (smnk) präpariert. Bei der Schichtzuweisung des Fund- occlusal pattern with the reduction of the paraconid stückes im Profil gab Herr Harald Oechsler (Wag- and the loss of the metaconid on the lower molars häusel) wertvolle Hinweise. Allen Genannten sei and the absence of carnassial cutting blade suggest a bestens gedankt. feeding mechanism unlike those of other creodonts and carnivorans (Lange-Badre & Böhme 2005), but close to that of mesonychids. Mesonychia, however, 110114_kaupia17.05.qxd.05.vrz 16.01.201118:05UhrSeite113 - ActaPalaeontologicaPolonica (MongolianErghilyin-Dzo People’s Republic). Ontwo Oligocene Hyaenodontidae from D. Dashzeweg, Africa: Cenozoic Mammals of (eds.): J. &Sanders W. Werdelin L. In: uemo Natural 4 History Museum of the American -Memoirs of middelEocene. Basin, Bridger Doebl Gesellschaft Naturforschende bergische überdieSencken- -Bericht Meeresand desMainzer Beckens. 1887 A. Andreae, Literature ar G. Mayr, S. Rietschel, & S.D. Peters, G., Mayr, W.D.Matthew, &Morlo, E. M. Lewis, L. &Trunkó, W. Munk, M., Leopold, W. Köppen, Forschungsinstitut Senckenberg, bei Rauenberg(Oberrheing Oligozän) derTongrube Bott-Eder Der »Rupelton« (Rupelium, T. &Schindler, A. Köthe, M.C., Grimm, K.I., Grimm, Folge. Neue Geologischen Mitteilungen desOberheinischen Vereins, -Jahresberichte und beiMalsch imKraichgau. rheingrabens A. &Hoppe, A. Foellmer, deFrance- Bulletin delaSociétégéologique duQuercy.) desPhosphorites fossile (Apterodon gaudryi P. Fischer, ExkursionD, Gesellschaft, - Exkursionsführer. - Carolinea, Pfalz, beiEschbach, Mittleren Oberrheingrabens von Mittel-Oligozän Landsäugetierresten immarinen des eln Heidelberg. Berlin, Ornithology -Journal of Germany. the Oligocene of ar G. Mayr, usa. umnbrs Science hummingbirds; ag-ar,B &Böhme,M. B. 2005. Lange-Badré, Berlin. Gruyter, brate Paleontology Verte- -Journal of Procellariformes). many (Aves: andBelgium Ger- from theOligocene of apekularfootmorphology with éet,Fac.-The Auk 116 France. Céreste, G. Mayr, Paris. - Annales dePaléontologie 91, (Germany). Espenhain anewEuropean Creodont Mammal from mp22 nov., sp. 75 543– F. , 1976 287 ( e osbr Clioms Coliidae) from Anewmousebird (Coliiformes: 2000. Anew 1999. 2004 nvriyo aionaPes Berkeley. Press, California University of 560; Note surunnouveau genre demammifère 1880. 46: ( – :Gudi e lmkne : derKlimakunde. Grundriß 1931): Tertiär (Baden-Pfalz). immittleren Rheingraben . 290 Karlsruhe. 13‒20; ) l ol oslrcr fmodern-type Old World fossilrecord of a): h anvr n netvr fthe Carnivora The andInsectivora of 1909. . Stuttgart. ; Ein neuesRaubtierausdemmitteloligocänen 22( Jahrestagung derPaläontologischen 46. Trogon 3 ): M. 304: 667– 25 Chapter 26 2010. . Die Randverwerfung Die desOber- 1993. raben, Deutschland). - Courier -Courier Deutschland). raben, 861– p alrh.D Karlsruhe. pp, : from themiddleOligocene of ered Ilinois, 676; Deerfield, ( 291 2 9 ), 237, 864; Washington d.c., ( – 2 427 567;New York. ): 263– 229– - Apterodon intermedius 434 , ( ( :Erster Nachweis 1990): 125 :Petrel-like birds 2002): akovle Florida, Jacksonville, ; Warshaw. 267; 253 – 133 – ›Creodonta‹. 8, rnfr .M. a. Frankfurt ; 1 –D25 rnfr .M. a. Frankfurt ; 280 388 brd. – 290 Karlsruhe. ; p De pp; 141 usa. 311– Paris. ; usa. , 2002. of 85 328 – 92; , ; - Biologist, G. Mayr, 182– 41 desGroßherzogtums Spezialkarte - Geologische Baden, Ornithology 148 Ornithology World hummingbird G. Mayr, schaften 91: -Naturwissen- Germany. songbird from theearlyOligocene of runelrfsi ie tla ora fZoology, -Italian Journal of Frauenweiler fossilsite. N. Micklich, Thürach,H. 27: -Mainzer geowissenschaftliche Mitteilungen, Befunde. derBasismikro- undmakrobotanischer Mainzer Beckens auf fürdenMittlerenstruktionen Rupelton (Unter-Oligozän) des (1998): Z. Kvacek, A., Bruch A. J., Pross, 269– Carnivora: New to theNatural Contributions of History (eds.): D. &Nagel, G. Gunnell, Morlo M., S. J. Mellett, &Manegold A. G. Mayr, Ornithology -Journal of preserved stomach content. Germany with fromGaviiformes) theearlyOligocene of G. Mayr, elr W. Weiler, H. Tobien, Gesellschaft Mainz, derRheinischen -Zeitschrift naturforschenden berg. der Tongrube Frauenweiler bei Wiesloch südlich von Heidel- ai. ücnrgois b. A, Abh., Basin.- Münchner geowiss. theUpper RhineGrabenandtheMainz the Paleogene of Espanol de Oceanographia Espanol deOceanographia -Publicationes EspecialesInstituto arevision. results of Preliminary Germany): Rupelian; weiler (Lower Oligocene, N.N &Parin, N. Micklich, Vertebrate Evolution Hyaenodon lypt(rb nefl) -Kaupia, clay pit(Grube Unterfeld). L. &Hildebrandt, N. Micklich, Modena. hog icn fLuai.In: Laurasia. through Miocene of carnivore guildsintheEocene analysisof – Ecomorphological Mittleren Carolinea, Rheingrabens.- drei tertiären Aufschlußkomplexen imRandbereich des W. &Munk, L. Trunkó, : 48 79– eln Heidelberg. Berlin, 186; ; Cambridge University Cambridge Press. 310; . Heidelberg. S.; Mainz. 92; e pcmn ftheearlyOligocene Old New specimens of 2007. Fossil hummingbirds intheOld World. 2005. 2004 52 ( (Mammalia, Creodonta). - Contributions to -Contributions Creodonta). (Mammalia, – 173 ( :TePsto fthe»GrandeCoupure« in ThePosition of 1987): :DieBedeutungderFischfunde imRupelton 1966): : 1904 ( ( 12– .Aprilseeo fanewfossilloon(Aves, skeleton Apartial of a. .Ploilg fNorth American Paleobiology of ). 1977 :Nwifraino h s an fthe New onthefishfaunaof information 1998): 177; , 105– 16;London. Erläuterungen zuBlatt Wiesloch (Nr. . 4 eln Heidelberg. Berlin, : 1:1– . - Journal - of Eurotrochilus. inexpectatus 111; 1 – Mainz. 37; 134 . Geologische Beobachtungen Geologische in 1998. eln Heidelberg. Berlin, ( 21 ;New York. :TheoldestEuropean fossil 2004): . , h sfuao Frauen- fishfaunaof The 1996. 129– owm .&Fica A. Friscia, & Goswami A. ( 148 :TheFrauenweiler 2005): 10: 56: 14 Madrid. ; ( : .Chapter 10 2010). 197– 9 113 – 28 Paläoklima-Rekon- – München. 202; Darmstadt. 118; Karlsruhe. ; 65 145 : 169– : 41 ). 184 ;

113 E. Frey,W. Munk, M. Böhme, M. Morlo & M. Hensel | First creodont carnivore from the Rupelian Clays of the former claypit Unterfeld at Rauenberg: Apterodon rauenbergensis n.sp. |