The Avifauna of the Miocene Fossil-Lagerstätte Sandelzhausen (Bavaria, Southern Germany)

22 Pages·2002·7.7 MB·English

by  Ursula B Göhlich

Zittcli.m.i
170
of V. Famlbusch (Municli). To date, these excavations havc InvestigationsbyHeissig(1997)onmammalbiostratigraphy
yielded approximately 30,000 identifiablc speciniens, oftheUpperFreshwaterMolasseinBavariashowedthefaunal
representingabout 170palaeozoologicalspccies(pcrs. comni. PositionofSandelzhausen in theearlv MiddleMiocene(MN5
V. Fahlbuscil), mostofthemvertebrates.Therichnessofthis = EuropeanMammalNeogeneUnitMN5,Badenian)withan
Continentalvertebratefaunaandtheextraordinarysystematic absoluteage ofaround 16 Ma. Several studies on the geology
diversity make Sandelzhausen an cxceptional site. The fauna and ondiffercnt faunal groups ofSandelzhausen ha\ealreadv
is dominated by vertebrates, although only represonted b\ been published (cited in F,\H1HUS( il & Lifbrhich 1996 and
isolated disarticulated remains. MoUuscs are rcpresented b\' ZiEGLhR 2000).
freshwaterandterrestrialgastropodsaswellasrarefreshwater
bivalves(Call 1972). Freshwaterostracods(Wrrr 1998),algal
gyrogonites,andfruitsofterrestrialangiosperms(Celtis, Gle-
1.2METHODS
ditsia)arealsorarecomponents.Withinthevertebrates,mam-
malsarethemostcommonfossils,followed by fishcs,amphi- Mostofthebirdremains,aswellasthousandsofinvertebrate
bians,andreptiles.Birdsaretherarestvertebratesfound,with andvertebratemicrofossils,wereobtained byscreen-washing
around 600 remains. The presencc ofbirds in the deposits of in thefield.The osteological terminology principally follows
Sandelzhausen has been known since the beginning of the Baumil et al. (1993) and occasionalK' Balimann (1969a, b);
diggings (Fahi.busch & Gau 1970: 393), butthevhave never nieasurements were taken after vdn hin Drusch (1976).
beendeterminedordescribed.Thisstudydealswiththeentire
bird material and presents the complete a\ifauna ofSandelz-
hausen.
Abbreviations
Ostceilogv - CMC: carpometacarpus, TT: tibiotarsus,
GEOLOGY
1.1 TMT: tarsometatarsus, dist.: distal, prox.: proximal, sin.:
Thefossiliferous limnotlu\latile deposits in Sandelzhausen sinister, dext.: dcxter, cran.: cranial, caud.: caudal.
reachamaximumof3metcrsinthickness.Ingencral,themarly Measuremcnts -GL:greatestlength,L:length,Bp:width
Sedimentsshowadecreaseofquartzpebblesinfrequencyand proximal, Bd: width distal, BTd: width Trochlea distal. Dp:
sizefromthebottomtothetop.In theIowerhalfofthesection diagonal proximal, Dd: diagonal distal, KC: smallestwidthof
there is a brown-colored "coaly" lignite layer of 2,5 - 5 cm corpus, Hp: height proximal, Hd: height distal, HpA: diago-
(fig. 7). Some horizons show a concentration of carbonate nalheightproximalwithapophysisflexorisinclusive,Td:depth
concretions. The fossils are conccntrated in and around the distal,Tp: depth proximal.
humous"coaly"laver,whilelarge-sizedfossilsaremoreoften
found in the deposits below the "coaly" laver. The deposits The matenal is housed in the Ba\ensche Staatssammlung
arguably cover a time span ranging from some hundred to fürPaläontologie und Geologie in Munich(BSP). Onlyafew
thousands ofyears (Schmid 2001). speciniensareheldattheStaatliches MuseumfürNaturkunde
in Stuttgart (SMNS).
Collections - BMNH: British Museum of Natural
History, London; BSP: Ba\erische Staatssammlung für Palä-
ontologie und Geologie, München; SAPM: Staatssammlung
fürAnthropologieundPaläoanatomie,München;FSL:Faculte
des Sciences de la Terre, Universite Claude Bernard Lyon 1,
Villeurbanne;ISEAK:InstytutSystematvki EwolucjiZwierz,
i
Krakow (Institute of Systematics and Evolution, Polish
Akademy of Sciences); LAC: Laboratoire d'Anatomie
comparce, Museum National d'Histoirc Naturelle, Paris;
LPVM: LaboratoiredePaleontologiedesVertebres,Universite
Montpellier; MALI: Magyar Ällami Földtani Intezet, Buda-
pest(HungarianGeologicalSurvey); ML Museumd'Histoire
Naturelle (Musee Guimet), Lyon; MNHN: Museum Natio-
nald'HistoireNaturelle,Paris;NMB: Naturhistorisches Mu-
50km seum Basel; RGM: Rijksmuseum van Geologie en Mineralo-
gie, Leiden (= Nationaal Natuurhistorisch Museum Leiden);
SMNS:StaatlichesMuseumfürNaturkunde,Stuttgart;UPM:
Miocene, Upper Freshwater Molasse Department des Sciences de laTerre, Marseille.
Country: A: Austria, D: Germanw F: France, 1: Italv, H:
Fiilgl.ust1r:atiGoenoogfrtahpeheixetpconssiitoinoonfotfhteheUpsipteerSaFnrdeeslhzwhaatuesrenMowliatshscscwhicmt.hiotuitc PHoulnagnadr,y,R:CZR:omCazmeac,hRSeLp:ubSlloivca,kEi:aS(psaeien,taMb.O6L):Moldavi.i,PL:
(^uatcrn.iryeover.
: 1 : 1
171
SYSTEMATIC PALAEONTOLOGY
2.
OrderAnseriformesWa(_,[.er, 1S3 Material and measurements (mm):
F.imil)' An.itidaeVlGORS, 1S25 BSP 1959 11
Suhtamily Dendrocheninae Livezey& Martin, 1988 HHuummeerruussdsienx.t,.,dipstr.oxe.ndhalf S822774 KKCC::55..42,,BBpd::(117.1)5
Genus Mwnctta LivEZE'i'& Martin, 1988 CMCsin.,prox. end 8380 Bp:9.1
CMC
dext.,prox. end 8292 Bp:9.1
CMC
sin.,dist.half 8301 Dd: 5.8
Mionetta blancbardi (Milne-Edwards, 1863) Radiussin.,prox. half 8288 Dp:4.3
(pl.l,figs. 1,2,3) Description:
Humerus: one proximal half (BSP 1959 II 8271) and one
Synon\-niv
distal end (BSP 1959 II 8274)ofa humerus are preserved.The
V 11896833 ADnednidrloicLihnechniibrldiiv-uhMairldinü{-ME]dInwea-rEdds':^.6\0r.d^, 1863)-Cmni-- crista bicipitalis is broken offon BSP 1959 II 8271.Thefossa
VM: 88-91,pl. 1,fig. 1 (with svnonymv). pthneeuompoetrnimcgipoiftathhesfisossshaalplnoewumaontdriccliopsietdah(snoins npenaerulmyahtoirci)zaonn-d
Locus tvpicus: St.-Gcr,ind-le-Pu\' (Allicr, Fr.incc). tal,alsoinventralview.Thefossapneumotricipitahsisenclosed
by a prominent edge, also at its cranial side. In ventral view
Stratum t\picuni: LowerMioccne,Aquit.inicn, MN2a. thecrusventralefossaeis rectilmear, nearU' horizontal,some-
Lectotypc: TT(MNHNAv8231)assigncdbyChenevai what ascending caudally; the tuberculum ventrale is barely
protruding distallv. On the caudal side the caput humeri is
(1983:88,91).
limiteddistalh'b\-adistinctmargin;initsventral halfthecaput
Original diagnosis: Milne-Edwards (1863: 160). IS somewhat undermined distally. The tuberculum dorsale is
roundish and prominent. The niargocaudalis flattcns in pro-
Extended diagnosis: Chene\ae(1983:90): «tibiotarsc ximal direction proximalK' before reaching the caput humeri
d'une taille intermediaire entre celle des especes aetuelles and the tuberculum dorsale. The suicus transversus extends
Dendrocygna javanica (denrocygne de Java) et D. viduata and deepens ventrallv. On the distal humerus, BSP 1959 II
(denrocygne veuf); surface articuiaire interne bien ereuscc, ä 8274,theepicondvlusventrahsandtheprocessusflexoriusare
bordposterieurtresdeveloppe;cretecnemienneexterneimpor- broken off. The imprcssion in the fossa musculi brachialis is
tante,cretefibulairelongue;gouttieretendineuselarge,äbord shallow,butdistinctandbroadensproximodorsally.Thefossa
interne tres fm et ä bord externe creuse d'une gouttiere du olecranibearsalongishhorizontalfurrow.Thesuicushumero-
peroneusprofundusprofonde;condyleinternetresfin(longu- tricipitalis is shallow and short.
eur= 77,7 mm, largcurde l'epiphysedistale = 9,1 mm).-
Carpometacarpus: Proximal end (BSP 1959 II 8380)with a
Stra11graph1ca1 and gcographicaI distribiition trochlea carpalis is somewhat extended proximodistally. The
France: St.-Gerand-le-Puy (Lower Miocene, MN2a, Mieni - Processus pisiformis is protrudingcramally. Processus exten-
Edwards 1863, Chenevae 1983); Germany: Ravolzhausen soriusissienderandlongwithaprocessusalularisprotruding
(LowerMiocene,MN1-2,Rheinhessen,Martini 1974),Weise- dorsally. The distal half (BSP 1959 II 8301) consists only of
nau(LowerMiocene,MNl,Lambrecht 1933);Czech Repub- the os metacarpale major; the os metacarpale minor is mis-
iic:Skiritz(LowerMiocene,MN3-4, LajMBRECEIT1933;Romania: sing. The cross-section of os metacarpale major is rounded
Credin ,a(UpperMiocene, Kessler 1992),Malusteni (Lower triangulär. The longitudinal tcndon furrow along the suicus
Phocene, MNl5, Kessler, 1992). interosseus narrows dorsallv. The shallow suicus tendineus
StartsatthemiddleleveloftheSymphysismetacarpalisdistalis
Remarks: Fraas(1870)described M. bhinchardi in Stein- and slightK' broadens as it extends proximocranially.
heim(MiddicMiocene,MN7),butvonAi\imc>n(1918)changed Radius:Theproximalhalf(BSP 1959II8288)witharoundish
this determination r.oAnas nsgoviensis. cotylahumeralis.Thetuberbicipitalisradialisisbluntandclose
Created as Arias blancbardi by Milne-Edwards in 1863, to the CLitvIa.The cross-section ofthe corpus is roundish.
Chenevae (1983) moved blancbardi to Dendrocben - Sub- Comparison and discussion: Withinthefourspecies
familyDendrocygninaeReichenbach, 1850-agenuscreated of anseriforms in Sandelzhausen Mionetta blancbardi is
by Miller (1944) fora humerusfrom the LowerMiocene of representedwiththemostspecimens(atleastfourindividuals).
South Dakota, USA. Based on osteological and phylogene- Themostdiagnosticfossilistheproximalhumerus,specimen-
tically significant characters of the Anseriformes, Livezey & BSP 1959 II 8271. It is characterized specifically by a shallow
Martin(1988)placedblancbardiintheirnewgenusMwnctta. andclosed(nonpneumatic)fossapneumotricipitahs;herewith
ELPirWeAvRinosu(s18o6s3t)e,olHogoiwcaalrddes(c1r9i6p4t)i,oCnhsewneerveaplro(v19i8d3e)d,bLyivMeziel:nye&- FitodlilfofwerisngfWrcom)OtlhfeegneenduesnA(>1i9a6s1a)annddtLhievAeznesye&nnMaaeritninge(n1e9r8a8l:.
Martin (1988). 198,207) thegenusAnas and the Anserinaeare characterized
byanopenpneumaticfossapneumotricipitahs.Althoughthe
: :
172
genusAythya is alsomarkodhyacloscdfossapneuniotricipi- Lcctotype: Ulna (MNHN Av 6428) figured by Milne-
talis(CnFNF\'Al 1987: 150),thcfossaisdeeperincomparison. Edwakds (1867-71: pl. 25, fig. 21-22), assigned bv Chenevae
Moreover tlie luimcrus of the Mioccnc spccies Aytbyd (1983:93).
chaiivimcisaiittlesmaller(Cheneval2000,tab. 1). Although Criginal diagnosis: Miene-Edwards(1867-71: 148ff).
clearly corresponding morphometrically wirb ,1/. hhinchanii
fromSt.-Gerand-ie-Puv,thepresentproximalhumerusdiffers Extcndcd diagnosis: Chenevai (1983: 93): "Cubitus
by one feature: thc cnclosure of thc fossa pneumotncipitalis d'une taille tres infcrieure ä ecUe de Dendrochcn blanchardi;
hasaprominentedgcallaroimd;thccraniodistalmarginclearly saillie du ligament articulaire anterieur peu marquec;
separates thc fossa from the shaft. Thc distal humerus (BSP iniprcssion du brachiahs anticus peu profonde, mais allongce
1959 II 8274) is morphomerricalU m accordance with M. le longdeladiaphyse; tubcrositccarpicnnc biencourbeevers
bhmchardi, but is also very similar to that ot Audi vclox. lecötemterne;dcpressionradialeinferieurcetroiteetprofonde
Comparisons show no distinct featurcs bctwccn the distal (longucur = 50,8 mm, largcur de rcpiph\sc proximale = 6,6
humerus cnd of M. blancljardi from St.-Gcrand-le-Pu\ and mm, largeurde rcpiph\sedistale = 5,3 mm).-
A. velox from Sansan. S11"a11graph1caI and gcographica1 distnbution
Bccause of the closcd fossa pncumotricipitalis and thc France: St.-Gerand-lc-Puy (Lowcr Mioccnc, MN2a, MlLNE-
relatively longerwing bones (in proportion to the leg boncs) Edwards 1863,Cheneval 1983);Germany:Weisen.iu(Lowcr
Mii.ne-Ed\xards(1863: 160)andLivezey&Martin(1988: 196) Miocene, MNl, Lambrecht 1933), Kastei Bruch (Upper
concluded that Mionctta blanchardi was moderately specia- Oligoccnc,Lambrecht 1933);CzcchRepublic:Skiritz(Lowcr
Miocene, MN3-4, Lambrecht 1933, Ml1kovsk> 2000a);
lized fordivint: rathcr than torflving.
Moldavia: Kishincv (Upper Miocene, MN9, Kessler 1992).
Material and measurements (mm):
Mwncitd ;;.i/aror(Miene-Edwards, 1867) BSP 1939 II
Coracoidsin.,prox. cnd 8269 KC:3.1
(pl. l,figs. 4, 5)
Coracoidsin.,prox.cnd 8982 KC:3.4
S\non\'m\' Humerussin.,prox.cnd
''V 1867 /1h<!5n<ir.!fo)-, nov.sp.-Miine-Edwards,vol. I: l4S,pl.23,
flg. 14-22.
V 1983 Dcndroiheti blambardi (Milnk-Edwards, 1867-71) -
CmNl\'Ai:93-94.pf I,flg.3 (withsvnonvmv).
Locus tvpicus: St.-Gcrand-Ic-Puv (Allier, France).
Stratum tvpicum: LowcrMioccnc,Aquitanicn,MN2a.
ZitU'liana 22, 2002 Plal
i
^m
J
Jf^
1a 1b
f
^'
^^
7b 3a
8c
1cm
12a 12b^ 13a 13b
GOHLICH,U. B.:The AvitaunaoftlieMiocene Fossil-Lat;crstättcSaiulclzhauscn (Bavaria, Southern Germany)
174
arecharactcrizedby.1roundcdofftriangular-sh.ipedanddcep Description and discussion: The specimens belong
cotyla scapularis, and by a shallow concavc facics articulans to an anseriform taxon of distinctiy larger size than the two
humcralis with a latcrally protrudmg margin. The iniprcssio species described above.
ligamentiacrocoracohumcralisisdistinctandlongishoval.The Ulna: Only represented bv two proximal ends. The
distaledgeoftheprocessusaerocoracoideus(tubereulumbra- olecranonprojeetsventrallyandsurpassestheroundishcotyla
chiale) medialK shows a little incision betwecn the facies ventralis ventrally. The cotyla dorsahs is protruding dorsally
articulares clavieularis. Whereas BSP 1959 II 8982 lies within and cramalK turns down distallv in form of a lip. DorsalK
themetriealvariabilityofM. ndtatorfromSt.-Gerand-Ie-Puy, beneath the processus cotylaris dorsalis lies a dent with the
BSP 1959 II 8269 is a bit smaller(own directcomparison). impressioscapulotricipitalis.Thetubereulumligamcnticolla-
UIna:Thepreserveddistalend represents atin\ ulnawith a teralis beneath the olecranon bears a distinct longish facies,
short but pointcd tuber earpale. A suicus mtereondylaris is thatdeepensatitsproximalhalf.Theimpressiobrachialis(only
present distoveiitrallv, but is absent distally. No incisura preserved on BSP 1959 II 8280) is longitudinal and narrow.
tendniosa ean be observed.
B\ the means of their size, these proximal ulnas probabK
Tibiotarsus: Only the distal fourth is preserved. The pons belongtothe Anserinae. Specimen BSP 1959 II 8280 is some-
supratendineus is damaged. Both condylus medialis and what smaller than BSP 1959 II 8281. However, comparisons
lateralis are sicnderand thereforethe incisura intcrcondvlaris with the measurements of some extant Anserinae (Bacher
is Wide. Alongthe preserved corpus,the suicus cxtensorius is 1967)showthatthedimensions(width prox. ofulna) ofboth
bounded niedially by a crest. The suicus niusculus fihularis, specimen are within the metrieal vanabilitv.
epicondylus medialis and lateralis and tubeiculum retinaculi
For most ofthe Necigene anserine species the ulnas are not
musculi fibularis are ver\- weak.
known. The proximal end of the ulna of Anscr oenmgensn
Humerus: Onlythepro.ximal end ispreserved.The interior (MiddleMiocene,Oeningen,Germany,vnNMeyer 1865)and
ot the fossa pneumotricipitalis is badly preserved, but seems ofAnscrobrantatan!bnkmi(UpperMiocene,Kishinev,Roma-
to be shallow. The opening of the fossa pneumotricipitalis is nia, KuR(x:hkin & Ganea 1972) aresomewhat larger; thatof
nearlyhorizontalinventral view.Thecaput humeri isslightly Cygiins cygniformis (Midtfle Miocene, Steinheim, Germanv)
damaged,caudally it is bordered distallv b\ adistinct margin IS distinctiy largen
and it is somewhat hollowed out distally in its ventral halt.
The tubereulum ventrale is barelv protruding distally. The Phalanx proximalis digiti ma|ons (Ph. dig. II 1): The com-
tubereulumdorsaleislongish,butweak.Thesuicustransversus plete preserved bone is long and siender. The pila cranialis is
craniallvflattened;dorsallyaweaksuicusrunsalongthewhole
e.xtends and deepens ventrally.
pila.Thecaudal wingis about thesamewidthalongthedistal
Comparison and discussion: AlldescribedClements -handdistallysurpassesthepilaalittlebit.Thewingshowsa
aredistinctiysmallerthanMionettablanchiardi,M. comobrina, longishandshallowconcavityontheventralsideanddorsally
A>husiuisdiiioisisandAythydchdiivirac.ThesmallsizedAnas therearetwoconcavitieswiththeproximal onemoreshallow
veloxfromSansanmetricallvcorrespondsinsomebones(e.g. than the distal one. Between them, there is a longitudinal
ulna dist.), or is only little larger (e. g. tibiotarsus dist.), but protuberance dose to the caudal end ofthe wing. The proxi-
theapparentshallow fossapneumotricipitalisofthe humerus mal end is triangulärwith a two-pointeddorsal end.Theven-
contradicts an allocation of these small sized anatids from
tralandcaudal marginoftheproximalendareweakU'concave.
Sandelzhausen to ÄJtas (see Woolfeneden 1961, Livezfy &
Martin 1988: 198,207).Theproximal humerus and thedistal Because of missing eomparative fossil material, all these
tibiotarsus from Sandelzhausen are a little smaller than those specimens are not determinable.
o{M.natatoriromSt-Gcrand-Ie-Puy(humenasprox.Bp: 13.6-
15.0mm,n=3;tibiotarsusdist.Bd:5.7-7.4mm,n=4,Chenevai Order Accipitriformcs Vllllioi, 1816
1983,tab. 1),butthelattcrrepresentonlyaveryfewspecimens.
The distal tibiotarsus from Skiritz (Czech Repubhc) is also a Three taxa of accipitriforms can be distinguished bv their
littlesmallerthanthose from St-Gerand-Ie-Puy (Bd: 5.3 mm, size, although they are not taxonomically determinable. For
Lambrecut 1933:358). morphometrical comparisons the following extant and fossil
(European Neogene) species have been considered:
Mionetta natatoy and Mionettd bLvicbctnii were formerK'
knownfromEarlyand LateMiocenedeposits(tab.6),thereby Extant: PaiiJion habaetus, Permi apivorus, Milvus milvus,
the specimens from Sandelzhausen are the first e\idence in Milvtis»iigydiii,Acdpitergentilis,Accipiternisus,Buteobuteo,
the MiddleMiocene. BiftcoLigopiii,Hieraactusfasciatus,Aquilachrysaetos,Aquild
helutca,Hebaeetusalbicilla,Hebaeetusleucocephalus,Aegypuis
monachns, Gypsjulvits, Circm cyanem, Circits aeriigniosiis,
Ansenformes gen. et sp. mdet. Falcopcregrinus,Falco sitbbiitco (thesestudied extant species
(pl. 2, figs. 6, 7) are housed in the SAPM).
Material and measurements (mm): Fossil: ButeopitsiUiis,AquiLipannatoides, A. delphmetisis,
Aquiliiviis depredator, A. corroyi, A. priHm. Hieraactus ed-
UlnaMii.,prox.end BSP82189059 11Bp: 12.6 wardi! (includmg Svn. Aquda nuiiuta), Habaeetiis pucator,
Ulnasin.,prox.end 8281 Bp: 14.1,Dp: 18.2 MilvHs deperditiis, Promibo incertus, Palaeohierax gervaim,
Phalanx prox.digiti majonsdext. 8379 L:33.7,GL:34.6,Bp: 8.7 Garganoaetiis jreudetithali, G. murivorus, Falco meduis.
175
PeLirgopappui magnus [thcse fossil species are studied eithcr BSP 1959II8320.Thelatterisrelativelydelicate.Thetrochleae
hx means ot tlicir original descnption or pcrsonally bv thc mctatarsiIIandIIIarethcsanicIcngthwhiletrochleametatarsi
.uitlior(torthclocalitics:St.-Gcrand-le-Pu\',Sansan,LaGrivc), IVISalittlcshorter. Indorsalvicwthctrochlea mctatarsi HIis
scctab. I]. vcr\'shghtlvobliquefromdistally lateral topro.ximalK' medi-
mm
al. Maximalwidthofthetrochleamctatarsi IIIis3.2 (BSP
1959 II 8320)and4.1 mm (BSP 1959 II 8997).Thefossameta-
FaniiK- AccipitridaeVieillüT, 1816 mm
tarsiIisslenderand8 inlength(BSP 1959II8320).Distally
Accipitridae gen. et sp. indet. A it extends to the level of the proximal end of the foramen
(tigs. 2, 3) vascularedistale.
Material and nicasurcnients (nmi): Tibiotarsus: The distal cnd of thc tibiotarsus is damaged
BSP 1959II caudally.Thetrochleacartilaginistibialiscaudallystrechesfar
TMTsin.distal halt 8320 KC:4.3, Bd: 10.8,Td: 7.0 proximally. The eondyli are relatively short. The canalis
TMTdext.distalcnd 8997 Bd 13,9,Td: 8.3 extensoriuspassesobliquclvfromdistallymedialtoproximally
TTsin.distalend 8308 Bd: 10.4,Td:(7) lateral. The oblique distal aperture of the canalis extensorius
PhalanxIV?terminalis(claw) 8329 Bp: 3.1,Hp:3.8,HpA:5.2 reachcs up to the medial edgeofthcdiaphysis. Theproximal
Phalanx I/II tcrmin.alis(claw) 8996 Bp 5.8,Hp:6.5, HpA: lO.I
aperture of the canalis extensorius is locatcd approximately
Descnption: midwidthofthediaphysis;ittapersoffdistallyandextendsto
Tarsometatarsus: One distal half of a tarsometatarsus sin. the levcl oftheproximal end ofthedistal aperture.
ancf one distal end of a tarsometatarsus dext. are preservccf. Phalanx IV? terminalis (claw): Thc small claw is ncarly
ThetarsometatarsusBSP 1959II8997issomewhatlargerthan complete,onK'theoutermosttipisbrokenoff.Thcapophvsis
Species
:
176
extensoris is long. The proximal articular facet is relatively HierdaetHsedwardsiisorientedmoreobliquelvindorsalview.
svmmetricalandpyi'iform.Theapophysisflexorisisalsolong Pdldcohierax gervdisn (St-Gerand-le-Puv, MN2, France) is
and cnds plantarclv to a pomted cdge. distincth' larger. Mdvm pcrdttiis (St-Gcrand-lc-Puv, MN2,
France) is only known by the holotype, a tarsomctatarsus
Phalanx I/ll tfiminali.s (claw): The claw is completc. It is withouttrochleaII. IncomparisionthetrochleaIIIisdistincth-
longandslenderanddistinctlv largerthanthcPh. IVtcrmliialis smallcn Promdio iiucrtiis (Chavroches, France) is similar in
describcd abovc. size, but differs by thc lack of the plantarely protruding
protuberancc on the trochlea IV. Pcldrgopdppm magnin and
Comparison and discussion: Bascd on rheir P. schlössen (Phosphorites ot Quercv, France, Eocene to
dimcnsions, thcseclemenrs niight bclongto thc same spccies. Ühgocene) as well as Gdrgduodctiis freitdentlhili and G.
Thcv rcprcscnt tlic smallcst of the thrcc recordcd taxa ot »lurivoriti (Gargano, Italv) are distincly larger.
accipitriformsinSandclzhausen.Thetworecordcdtarsometa-
tarsi differinsize. But studicson extantaccipitriformes show UntortunatelvthercaresomeNeogeneaccipitndtaxawhere
thatthesizeotthetarsometatarsican var\' upto25"/o (conccr- both the tarsomctatarsus and tibiotarsus are not known - likc
ningthedistalwidth)e.g.inAcctpüergracilis,whichisknown the fossil buzzard Butco pusilltis (carpometacarpus from La
MN
tohavethelargestdimorphisniinsize(SrHMil)T-BuR<;FR 1982: Grivc, 7+8, B.\i imann 1969a) orthefossil eagleHalidc-
116). Sotheditfercnce in sizeofthetwodescribcd tarsometa- ctiis piscdtor (carpometacarpus and phalangcs from Sansan,
tarsi,amountingto22'/o,canassumedtobewithmtheVariation Mii.Ni-EuwARMs 1867-71, Cheneval2000). After Balimann
ofone species. Bevond that, the two tarsometatarsi are mor- (1973: 1Of),thetarsomctatarsusofbuzzardsandeaglescanonlv
phologically identical. Thc tibiotarsus and tarsomctatarsus bedistinguishedbvcharactersofthcproximal cnd,butwhich
differfrom thcfamilv Falconidac bv the lackofan additional is not yet represented in Sandclzhausen. So with the present
second distal opening of the canalis extensonus on the osteologicalknowledgeoffossilaccipitrids,adefinitivesyste-
tibiotarsus and bv thc more plantar position of the trochlea matic determination ofthcse bones is not possible.
metatarsi IVancithemorcmedialoncntationoftheprocessus
of the trochlea metatarsi II on the tarsomctatarsus (in distal Thcclawsarerefcired tothis taxononaccountoftheirsize;
view).ComparedwithextanttaxaotAccipitridacandbvmcans thcvaretoosnialltobclongtothebothlargertaxaofaccipitri-
of morphology, thc tarsomctatarsus and the tibiotarsus scem forms from Sandclzhausen. Metrically the phalanges arevcrv
to bedosest to the buzzard Butt'o butco. similartotheclawsofdigit IVand I/IIofextant Butcobutco.
The Tertiary European species likc Aquilaviis dcpredator,
A. prisciis, Aqiiila delphmeiisis, A. pcnnatoidcs, Palacohicnix Accipitriformes gen. etsp. indet. B
gervaisii,GargMiodctmfrcitdcutlidli,G. miirivorus,andPcLu- (fig. 5)
gopappas magniis are distinctlv larger (see tab. 1). The distal Material and mcasurements (mm):
tarsometatarsiofbothAi/mlapannatoidesandA. delphinensu
fderlopmhiLnacnGsrisivhcas(MaNm7u+cSh,bFrroaandccer)sahraeftnotthaknncoawnnb.eHaoswseuvmeerd,f^o4r. PPlhuallaannxxI1 I1 ddcexxttB..SPS8331747Sh79 1LL1::2233..53,,BBpp::99..57,, HHpp;: 55..15,, KKCC:: 55..03,, BBTTdd::65..07
thc present specimcns. AqmLivits dcpredator and A. pnsciis
(bothfromSt.-Gerand-lc-Puy,MN2,France)ditferb\-alarger Description
distaltarsomctatarsus.Aqndaviiscorroyi(from the Palet>gene
ofQuercy, France) is similar in size, but isdistinguished bv a Phalanx 1 1: Both specimcns are complete. The proximal
shortencd trochlea IV (m dorsal view) and a lacking plantar cnd is dorsoplantarelv flattened and shows a medial indenta-
tion. Thc plantar side ofthe shaft is weakly concavc. The di-
protuberanccon trochlea II (in medial view).
stal trochlea is tiattcncd proximalK'.
Piimdio inccrtus (formerly in Milviis) from the Lower Comparison and cfiscussion: These phalanges re-
MiocencofSt-Gcrand-le-Puy(France)ismetricallysimilarto present the middle-sizcd of the three recordcd accipitriform
thedistal cndofthctarsomctatarsusfrom Sandclzhausen, but taxa. Asmentioncdbetöre,taxonA iscomparablcinsizewith
indistalviewthemedial processusofthetrochleametatarsalis extant Butco butco and taxon C is in the size ränge of the
II isshorterand thcplantarprocessusofthetrochleametatar- GoldenEagle(Aquddchrysactos),whichwillbeshownbelow.
salis IV IS not present m P. nicertns. Compared with thcse spccies and considering the sexual
Inthedistal halfofthctarsomctatarsusofMdvmdcpcrdittn dtoimtohrephiacscniip,ittrhiedphtaalxaonngeAsIaInadretdoefoinsimtailvellvtotobocbliogntgobteolotnhge
from thc Lower Miocenc of St-Gerand-le-Puy (France) the
accipitnd taxon C.
tossa metatarsi I is somewhat elongatcd proximally, thercby
longer. Additionally, in contrast to the Sandclzhausen spcci- Therc are onl\ a few taxonomically determined fossil pha-
men,theprocessusofthetrochleametatarsalisII indorsalview lanx I 1. Three specimcns are known from Sansan and deter-
surpassesthctrochleadistallyand indistal view itarisesfrom mined all to bclong toHicrdcetusedwdrdsi(Cheneval 2000:
the trochlea metatarsalis II by a wcll-markcd angle. 341, tab. 2). According to own studics the Variation in sizeof
thcse phalangcs scems to be too large to refcr them all to the
Phcrddetiiicdwdrdii(S3.n^M-\,MN6,France)issimilarinsize, samespecies.ThephalanxI 1 frc^mSandclzhausencorrcspond
but ditters in morphology: thc trochlea metatarsalis III of to thc two smallcrspecimcns trom Sansan.
177
Fig. 2: Accipitriformes gcii. et sp. indct. taxoii A, tarsomctAtarsus ^
sin.dist.ilhalf(BSP 1959II8320).a:dorsal,b;lateral,c:plan-
tar,d: medial,e:distal.
Fig. 3: Accipitriformesgen.etsp.indet.taxonA,tibiotarsussin.di-
stalend(BSP 1959II 8308).a:cranial,b:caudal,c:distal. Fig. 5: Accipitriformes gen. et sp. indet. taxon B, phalanx I 1 dext.
Fig. 4: Accipitriformes gen. et sp. indct. taxon C, plialanx I/II (BSP 1959II8377).a:dorsal,b:plantar,c:proximal.
terminalis(claw)(BSP1959II8327).a;lateral/medial,b:pro- Fig. 6: Accipitriformesgen.etsp.indet.taxonC,PhalanxIII3(BSP
ximal. 1959 II 8378).a:dorsal,b:plantar,c:proximal.
:
178
Accipitriformes gen. et sp. indct. C wards 1867-71:pl. 183,fig.26-29)islargerandsomewhatmore
(figs. 4, 6) slender. Very close in size and morphology are the fossils of
Halmetiispiscator from Sansan (Cheneval 2000: 337ff). But
Material and measurements (mm):
the scanty material allows no allocation.
BSP 1959II
Phalanx I/Il tcrminJis(claw) 8326 Bp: S.6,Hp:9.8,HpA: IS.3
Phalanx I/ntcrmin.ilis(claw) 8327 Bp:9.1,Hp: 10.5,HpA: 15.1
Phalanx III 3 8378 GL:26.0,Bp:6.6,Hp:6.2, Order GalliformesTemminc;k, 1820
BTiI:6.4
Family PhasianidaeVKiDRs, 1825
Description:
The galliforms are represented by 3 taxa of Phasianidae in
Phalanx I/Il tciminalis (claw): The tvvo claws with liroken theavifaunaofSandelzhausen.Withinthegalliformspartridges
offtipsbelongto thedigit I or II. Bothspccimcn showa long are in general represented morc abundantly in the European
apoph)'sis extensoris and apophysis flexons. Both the apo- Miocene than pheasants (Cheneval 2000: 343), as is the case
physis extensoris and flexoris ofBSP 1959 11 8327 are a little in Sandelzhausen. Most ofthe galliform reniains of Sandelz-
longerthanthoseofBSP 1959 11 S326,especiallv indorsaland hausen belongtoPaLieortyxintermedia followed in quantity
plantarview. Theproxmial two-parted articularfacet is pvn- by Miophiniaina altus\ only two fragments represent a tiny
form and svmmetncal.
Phasianidaeincfet..P^Ljeortyxmtermeduihasbeenknownfrom
Phalanx 111 3: Inlateral viewthephalanx is bentproxiniallv olderdeposits and can nowbe confirmed in MN 5 (tab. 6).
convex. The proximal two-parted articular facet is relatively
symmetrical. The plantar side of the corpus is concaxe. The
Genus Miophdsutnns LAMBREt:EiT, 1933
distal trochlcaisdorsoplantarely flattened.
Miophdsianiisdhus (Miene-Edwards, 1869)
Comparison and discussion: These phalangeal
(pl.2, figs. 1-4)
reniains represent the large sized accipitrid taxon. In lateral
view the claws show a characteristic apophysis extensoris as Synonymy:
well as a long protruding slender apophysis flexoris. Both '' V 1869 PhdShvniutltiii,nov.sp.-MiLNi-Edwards,vol.2:239-241,
featuresareknownfrombuzzards,eaglesandhawks.Vultures, pl. 131,flg.27-36.
however, generally do not show a clear apophysis extensoris V 2000 A/zo/i/a!»;.»;;/).i/ri/s(Milne-Edwards,1869-71)-CiiENEyAL:
(Bali.mann 1973: 12).Theclawsaredistinctlylargerthanthose 351-352,figs.9-11 (withsynonymy).
ot Hicraaetiis edwanisi from the Middle Miocene of Sansan Lectotype: tarsometatarsusdext.,prox. end(MNHNSa
(France,seeMilne-Edwards 1867-71,Cheneval2000).How- 1221),lectotypefixationbyCheneval(2000:35If),figuredin
ever, they are morphometrically similar to those from St- Milne-Edwards (1869-71: pl. 131, fig. 27-29).
Gerand-le-Puy(France,LowerMiocene,seeMune-Euwards Locus typicus: Sansan (Gers, France).
d1e8t6e7r-7m1i:nepdl.. 183, figs. 17-21), which are not taxonomically Stratum typicum: Middle Miocene, Astarac, MN 6.
Original diagnosis: Milne-Edwards 869-1971
In comparison with extant accipitritorms the phalanx 111 3 (1
is very close in size and morphology to that of the Golden 239ff): «L'uned'elles[GallinaccsdeSansan]paraittres-voisine
Hagle (Aquila chrysactos). Compared with the few known des Faisans,etäraisondesesdimensions,je l'aidesigneesous
suitablefossils,thephalanges1113oiHieraeetusedwardsifrom le nom de Phdiidnmdltits.»
Sansan are smaller (see Cheneval 2000: tab. 2), and a single «La portion superieure du metatarse de ce Faisan presente
undetermined specimen from St-Gerand-le-Puy (Miene-Ed- en effet les characteres propres aux oiseaux dont le genre
Plate 2
Figures 1-4in natural size,figures5-10 indoublesize.
Miophasiiintisdltiis(Milne-Edwards, 1869)
Fig. 1: Fcmursin.;a:cranial,h: caudalc: proximal,d:distal(BSP 1959 II 8382).
Fig. 2: Tarsometatarsusdext.;a:dorsal,b: plantar,c:distal(BSP 1959 II2207).
Fig. 3: Ulnadext.distalend;a:dorsal,b:ventral(BSP 1959 II 8980).
Fig. 4: Tibiotarsussin.;a:cranial,h: lateral,c:caudal,d:proximal,c:distal(BSP 1939 II 8381).
Palacorlyxintcrnifilid Balimann, 1969
Fig. 5: Tarsometatarsussin.,distalend;a:dorsal,b:plantar(SMNS54006/1).
Fig. 6: Humerussin.,distalend;a:cranial,b:caudal(BSP 1959II 8273).
Fig. 7: Phalanxdig. II 1 dext.;a:dorsal,b:ventral (BSP 1959II 8331).
Fig. 8: Coracoidsin.;dorsal(BSP 1959 II 8270).
Fig. 9: Tibiotarsusdext.,distalend;a:cranial,b:caudal (BSP 1959 II 8307).
Phasianidae indct.
Flg.10: Carpometacarpusdext.,distal2/3;dorsal(BSP 1959 II 8291).