reappraisal of testudo antiqua

19 Pages·2014·2.1 MB·English

JournalofPaleontology,88(5),2014,p.948–966
Copyright!2014,ThePaleontologicalSociety
0022-3360/14/0088-0948$03.00
DOI:10.1666/13-036
REAPPRAISAL OF TESTUDO ANTIQUA (TESTUDINES, TESTUDINIDAE)
¨
FROM THE MIOCENE OF HOHENHOWEN, GERMANY
JOSEPH A. CORSINI,1 MADELAINE BO¨HME,2,3 AND WALTER G. JOYCE2,4
1DepartmentofBiology,EasternOregonUniversity,LaGrande,OR97850,USA,,[email protected].;2DepartmentofGeosciences,
UniversityofTu¨bingen,Ho¨lderlinstr.12,72074Tu¨bingen,Germany;3SenckenbergCenterforHumanEvolutionandPalaeoecology,
Ho¨lderlinstr.12,72074Tu¨bingen,Germany;4YalePeabodyMuseumofNaturalHistory,170WhitneyAvenue,NewHaven,CT06511,USA
ABSTRACT—Testudo antiqua is one of the few fossil turtle names to have survived the past 200 years of taxonomic
reshufflingwithitsoriginalgenusandspecificepithetintact.Theninecurrentlyknownspecimenswerecollectedfromthe
middleMioceneHohenho¨wenlocalityinsouthernGermany.BecausetheavailableHohenho¨wenmaterialwasneverfully
described,weherecompletelydocumentallknownspecimens.Itisunclearwhichofthesespecimensformedtheoriginal
T. antiqua type series, so we herein selected the best preserved representative as the neotype. A phylogenetic analysis
places T. antiqua in a basal polytomy within the clade Testudo, indicating that T. antiqua may represent the ancestral
morphologyofTestudo.Aswithanumberofotherpublishedstudies,ourswasunabletoresolverelationshipsbetweenthe
three extant Testudo lineages (the hermanni-group, the graeca/kleinmanni/marginata group, and the horsfieldii-group).
Finally,withaviewtowardlocatingmoreturtlesandinordertobetterunderstandthegeologicalandecologicalcontextof
these tortoises, we visited Hohenho¨wen several times to search for the original collection sites, but we were unable to
locate the original fossil quarries described in the literature.
INTRODUCTION taxonomic revision in the past four decades, and considerable
IN THE 250 years since Linnaeus placed all known turtles into controversy has been associated with the nomenclature of these
Testudo, the genus has been reduced to only five extant turtles (e.g., Khozatsky and Mlynarski, 1966; van der Kuyl et
species (e.g., Joyce et al., 2004; Fritz and Havas, 2007; Rhodin al., 2002; Pera¨la¨, 2002; Bour, 2004: Parham et al., 2006a,
et al., 2008) and a handful of closely related fossil forms (e.g., 2006b; Lapparent de Broin et al., 2006a, 2006b, 2006c; Fritz et
Lapparent de Broin et al., 2006a, 2006b, 2006c). Perhaps the al.,2007;FritzandBininda-Emonds,2007).Itisclearfromboth
best preserved among these fossil forms is Testudo antiqua molecular and morphological studies that all of the close
Bronn,1831,oneofthefirstfossilturtleseverdescribed(Kuhn, relatives of T. graeca (i.e., T. marginata Schoepff, 1792, T.
1964). Through coincidence of its close relationship with the hermanni Gmelin, 1789, T. horsfieldii Gray, 1844, and T.
typespeciesofTestudo,T.graecaLinnaeus,1758,itisalsoone kleinmanni Lortet, 1883) form a monophyletic clade, and some
of the few fossil turtles named in the early nineteenth century molecular studies have provided support for inclusion of
that has not been referred to a new genus during the past two Indotestudo and Malacochersus in this clade (see phylogenetic
centuriesofsplittingandrenaming.TheoriginalspecimensofT. analysis below). It is our opinion that taxonomic endeavors
antiqua were collected from the Hohenho¨wen site in south- should aim at stabilization of names within well-defined
westernGermany,avertebratelocalitywhichisnowconsidered monophyletic clades (e.g., Parham 2006a, Joyce et al., 2004),
to be of middle Miocene age (ca. 13 Ma, see below). Testudo so in this work we follow Fritz and Bininda-Emonds (2007) in
antiquawasinitiallydescribedbyBronnin1831,whocompared retaining the original taxonomic designation for all of these
it to extant T. graeca and Chelonoidis denticulata Linnaeus tortoiseswithinTestudo.ThisincludesT.horsfieldii,whichwas
1766 (then T. tabulata). The illustrations in that original placed into the genus Agrionemys by Khozatsky and Mlynarski
description are highly idealized, so the actual features of the (1966),T.hermanni,whichwasplacedinthegenusEurotestudo
originally described specimens are impossible to determine. by Lapparent de Broin et al. (2006a), and T. marginata and T.
Testudo antiqua was re-described by von Meyer in 1865, who kleinmanni, which were placed in the genus Chersus by Gmira
produced more accurate illustrations that were also idealized in (1993).
some aspects. More recently, Schleich (1981) included T. Determination of the exact phylogenetic position of T.
antiqua in a large morphometric study of fossil turtles from antiquahasbeenhamperedbythelackofacompleteillustration
Germany, partially illustrating four specimens, photographing and description of all the known material, so it is not known
others, and finally designating a lectotype from the alleged whetherT.antiquaisancestraltoorpartofthemodernTestudo
syntypeseries.EightspecimensofT.antiquaarecurrentlyheld clade.Testudoantiquawasincludedinalargestudy(Lapparent
in German collections (FFSM, SMNS, UFGC). A ninth de Broin et al., 2006b, 2006c) that elucidated the relationships
specimen from Hohenho¨wen is housed in a French collection between fossil and extant Testudo, and the authors concluded
(MT),andwaspartiallyphotographed,thoughnotillustrated,by that T. antiqua lies along the T. hermanni lineage. However,
Broin(1977).Itisapparentfromthehistoricaldescriptionsthat they did not have direct access to any of the specimens held in
other specimens were at one point available but their Germancollectionorcompleteillustrationsoftheentiresyntype
whereabouts are now unknown. series,andthescoringofT.antiquawasthereforeinaccuratefor
Testudo antiqua belongs to a widespread group of closely severalcharacters.Becauseoftheaforementionedquestionsand
related tortoises than have inhabited Europe, Asia, and North phylogeneticambiguities,wehereinfullyillustrateanddescribe
Africa since the Miocene. This group has undergone major allofknownT.antiquamaterialfromHohenho¨wen.Withwell-
948
CORSINI ET AL.—TESTUDO ANTIQUA FROM THE MIOCENE OF HOHENHO¨WEN 949
FIGURE1—1,theHohenho¨wensiteislocatedinBaden-Wu¨rttemberg,nearthesouth-westernGermanvillageofEngen;2,measuredsectionofHohenho¨wen
outcrop.TheDeckentuff,whichoverlaysthefossiliferousgypsum,hasbeendatedat12.5–13Ma.
described material, we then cladistically explore the phyloge- Geology.—The Hohenho¨wen (or Hohenheven) locality in
neticrelationshipsofT.antiquausingamodifiedversionofthe southwestern Germany (Fig. 1.1) was first described as a
character matrix reported by Lapparent de Broin et al. (2006b, freshwatergypsum(Su¨sswasser-Gips)intheearly1800s(Bronn,
2006c).Theresultingtreeisthencomparedtorecentmolecular 1831). The site is located on the northeast flank of the
trees, and the possibility that T. antiqua displays the ancestral Hohenho¨wenpromontorynearthevillageofEngenintheHegau
morphology of modern Testudo is raised. region of southwestern Baden-Wu¨rttemberg, Germany
(Schreiner, 1983, 1992; Fig. 1.1). The fossiliferous layers were
GEOLOGICAL AND TAPHONOMIC SETTING apparently first collected at the beginning of the early 1800s,
Institutional abbreviations.—FFSM, Fu¨rstlich Fu¨rstenber- perhaps exposed by a massive landslide involving the entire
gisches Sammlung Donaueschingen, Donaueschingen, Germany; eastern flank of the massif in 1817. The Hohenho¨wen hill is the
MT,MuseumdeToulouse,Toulouse,France;SMF,Forschungs- secondhighestelevation(846mmabovesea-level)oftheHegau.
institut und Naturmuseum Senckenberg, Frankfurt am Main, The top of this hill is comprised of a basalt chimney, which
Germany; SMNS, Staatliches Museum fu¨r Naturkunde Stuttgart, penetratesthe250mthickJu¨ngererJuranagelfluh(JJ)unit(Fig.
Stuttgart, Germany; UFGC, University of Freiburg, Geological 1.2; Schreiner, 1992). The JJ unit is a clayey to silty marl
Collections, Freiburg, Germany. containing pebbles of Mesozoic limestones, sandstones, and the
950 JOURNAL OF PALEONTOLOGY, V. 88, NO. 5, 2014
Variscan basement. It is part of the Upper Freshwater Molasse scavengingbyrodents(viagnawing)orlargeranimalsthatleave
along the northern margin of the western North Alpine Foreland tooth marks on the shell bone is often observed on many
Basin and is interpreted as the erosional detritus deposited into specimens (forexamplethe White River tortoises[Corsiniet al.,
alluvial fans by the rising Black Forest Mountains (Schreiner, 2006]or thelate middleMiocene faunafrom Gratkorn [Grosset
1992).TowardthetopoftheJJatHohenho¨wen,bedsofmassive al., 2011,fig.6a]). Together,theabsence of non-shellbonesand
pedogenic gypsum have yielded pockets of fossil vertebrates absence of gnaw marks suggests an ecosystem devoid of small
(according toWalchner,1851,p.983,shellsof10to12Testudo mammals able to scavenge calcium but containing scavengers,
antiqua individuals were found in close proximity). The gypsum perhaps birds, able to remove the non-shell bones.
bedsareoverlainonthesouthwestflankofHohenho¨wenat730m
by 10 m volcanic tuffs (Fig. 1.2), which, according to Schreiner SYSTEMATIC PALEONTOLOGY
(1992), can be correlated with the Deckentuff (700 m) at the TESTUDINES Linnaeus, 1758
nearby Hohenstoffeln locality. The latter tuff has been dated by CRYPTODIRA Cope, 1868
K-Ar method to 12.5 Ma (sanidine; Lippoldt et al., 1963) and to TESTUDINIDAE Gray, 1825
12.8–13.0 Ma (hornblende; Schreiner, 1966, 1983). These ages TESTUDO Linnaeus, 1758
compare well with biochronological data retrieved from mam-
malianfossilsfoundtogetherwithT.antiquaattheHohenho¨wen Type species.—Testudo graeca Linnaeus, 1758.
site.AccordingtoStehlin(1926)andTobien(1957)thefollowing
mammalian species are recorded: Lagospis verus, Gomphothe- TESTUDO ANTIQUA Bronn, 1831
rium sp., Anchitherium aurelianense, Euprox furcatus, and Neotype.—Given that we were not able to identify a single
?Micromeryx flourensianus. This association resembles the individualfromBronn’s(1831)originalsyntypeseries,weherein
well-known JJ locality Anwil (Switzerland, 75 km southwest of designatethebest-preservedspecimen,MTPAL2012.0.10,asthe
Hohenho¨wen), which contains all of the aforementioned animals neotype, and abandon the lectotype designation by Schleich
except Gomphotherium sp. (Engesser, 1972). Anwil is biostrati- (1981) (see Remarks below).
graphically dated to about 13.3 Ma (Ka¨lin and Kempf, 2009). Diagnosis.—Testudo antiqua is diagnosed as a member of
Thus, both radiometric dating and mammalian biochronology Testudinidae by the presence of a thickened epiplastral lip,
support an absolute age of the Hohenho¨wen fossils of about 13 coincidence of the costal/peripheral suture with the pleural/
Ma (MN8, middle Miocene). It is worth noting that we revisited marginal sulcus, wedged-shaped costals, and high peripherals.
Hohenho¨wen several times and were unable to relocate the TestudoantiquacanbeplacedwithinTestudobythepresenceof
original quarry, which appears to have been covered by more an elongatecervical scute, a narrow nuchal notch defined by the
recent landslides and is now overgrown by forest. This is a first marginals, a quadrangular pygal that is wide anteriorly and
poignant reminder that fossil localities do not necessarily last narrow posteriorly, a humeral-pectoral sulcus that does not cross
forever. entoplastron,anepiplastralexcavationthatpenetratesnearlyhalf
Taphonomyandfaunalassociations.—Testudoantiquaappears of the thickness of the anterior portion of the epiplastron, two
to have been part of a vibrant ecosystem evidenced by diverse suprapygals, and a suprapygal 1 that embraces the half-moon
terrestrial gastropods (Seemann, 1930) and fragmentary post- shaped suprapygal 2. Testudo antiqua differs from T. graeca, T.
cranialremainsofcarnivorans,elephants,horses,deer,andpikas marginata,andT.kleinmanniandtheirfossilrelativesinlacking
(Stehlin, 1926, Tobien, 1957). All of these fossils are housed at aposteriorplastralhinge,andfromT.horsfieldiianditsrelatives
FFSMandareinneedofrevision.Thehostsedimentisamassive bybeinghigh-domed.ThevertebralseriesdiffersfromT.graeca
pedogenic gypsum bed (i.e., Su¨wassergips) that contains red and inbeingsomewhatnarrowerthanthevertebralseriesofT.graeca
yellow-mottled clays and marls (Rote Letten of von Althaus, but not as narrow as that observed in T. hermanni.
1832). This type of rock indicates seasonal arid climate Type locality.—Hohenho¨wen, Engen, Baden-Wu¨rttemberg,
conditions during deposition with mean annual precipitation Germany. Pedogenic gypsum, middle Miocene, MN8 (~13 Ma,
below300mm(Retallack,1994).Anolddescriptionofthesiteby also see Geological Setting).
Walchner(1851)suggeststhatallofthefossilswererecoveredin Referred material.—FFSM 3446.1, FFSM 3446.2, FFSM
close proximity to each other, although the original locality and 3446.3, FFSM 3446.4, SMNS 4450, SMNS 51467, SMNS
collection descriptions are not clearly described. Interestingly, in 51469, UFGC 9.
contrasttothemammals,nearlyalloftheturtlesareinrelatively Remarks.—The original description of T. antiqua is based on
high state of articulation, suggesting that burial was rapid. shell material from three to four specimens collected at
ControlledstudiesbyBrandetal.(2003),aswellasobservations Hohenho¨wen, but Bronn’s (1831) descriptions and illustrations
byDodd(1995)andCorsiniandChamberlain(2008),suggestthat areinsufficientforidentificationoftheindividualsinthatoriginal
turtles completely submerged in water disarticulate within two syntype series within the available material. Von Meyer (1865)
years,whileinsemiaridtoaridconditionsfreshwaterturtleshells clearlyillustratedFFSM3446.1,perhapsalsoUFGC9,andstated
exposed on the surface completely disarticulate within three to thatBronnexaminedthosetwoinhisoriginalwork,butwewere
fouryears(Dodd,1995;Brandetal.,2003).Soitislikelythatthe unabletoindependentlyverifythisassertion.Schleich(1981)felt
Hohenho¨wen tortoises were buried quickly, within months of compelled to designate SMNS 4450, a poorly prepared and
death. Interestingly, no non-shellboneswere recovered withany extremely deformed individual, as the lectotype, probably
of the turtles, and we observed only one possible non-shell bone becauseitwasreadilyavailableattheStuttgartMuseum(SMNS).
fragment among all of the steinkerns. Our experience with other However,giventhatwefoundnoevidencethatSMNS4450was
tortoiseassemblages(Corsinietal.,2006)suggeststhatnon-shell indeedpartofthesyntypeseries,thedesignationofalectotypeis
elementsareobservableinone-halftothree-quartersoftheturtles dubioustous,andwealsoquestiontheassignmentoftypestatus
observed in the field, so with nine specimens there is a high toaspecimenwithsomanymissingcharacters.Karl(2013)more
probability that if non-shell elements are present they should be recently argued that the specimen housed at the University of
observableinatleastseveralcases.Thissuggeststhatmanyofthe FreiburgGeological Museum (UFGC9) shouldbethe lectotype,
non-shell bones were removed from the carcass by scavengers. basedmainlyuponthefactthatvonMeyer(1865)wasthoughtto
Apart from the missing non-shell bones, there is no overt have viewed that specimen. However, he makes no clear
evidence of scavenging or other post-mortem alteration (bite or connection between that specimen and the specimens viewed by
gnawmarks).Thisissurprising,becauseinotherlocationswhere Bronn in 1806. The designation of a lectotype is therefore not
high concentrations of tortoises are recovered, calcium valid, as lectotypes must demonstrably be part of the original
CORSINI ET AL.—TESTUDO ANTIQUA FROM THE MIOCENE OF HOHENHO¨WEN 951
syntypeseries.Wethereforeconcludethataneotypedesignation the pygal, and right peripherals 8, 9, 10, and 11. The distal
is appropriate, and designate MT PAL.2012.0.10 as the neotype portions of right peripherals 4, 5, 6, and 7 are missing as well.
because: 1) it originates from the type locality; 2) it is the best- Also missing are the lower sections of all of the left costals, the
preservedspecimen;and3)itiseasilyaccessibleinalargepublic leftedgeofthepygal,andthelowerleftterminusofsuprapygal2.
collection. We finally note that one specimen at FFSM is also Ontherightalargesectioncontainingthedistalhalftotwo-thirds
quite well preserved, but we omitted this specimen from of costals 2, 3, and 4 is absent, and the lower half of costal 5
consideration, because it is not housed at a regular, public shows damage. Approximately 75 percent of the plastron is
museum. present,althoughtheepiplastron,partoftheentoplastron,andthe
anterior portions of the right hyoplastron are still encased in
DESCRIPTION matrix. Missing from ventral view are large portions of the left
FFSM 3446.1.—Specimen FFSM 3446.1 is an approximately and right hyo- and hypoplastra, and the entire xiphiplastra.
95 percent complete shell with some dorsal and lateral However, the xiphiplastra and the right portion of the right
deformation toward the left (Fig. 2). No cranial or post-cranial hypoplastron are present as a large detached fragment (not
elements are present. The characteristic (for Hohenho¨wen) red- figured), but can only be viewed from the visceral side because
orange mottling of the bone surface is present. Missing from the the ventral surface is encased in matrix.
carapaceistheouterrightportionofperipheral1,asmallpieceof SMNS 4450.—This specimen (proposed lectotype of Schleich
the border of peripheral 2, the posterior half of peripheral 11, a 1981)isanapproximately85percentcompleteshellthathasbeen
smallpiecefromthecentralportionofthedistalendofperipheral significantly deformed, primarily through dorsal-ventral com-
10,thepygalbone,partsofcostals7and8,andasmallpieceof pression(Fig.6).Nocranialorpost-cranialelementsarepresent.
costal6.Thesuturebetweenneural7and8isnotvisible,andthe Theshellboneshavethered-orangemottlingthatischaracteristic
junctions of left costals 7 and 8 with neurals 7 and 8 are ofHohenho¨wenspecimens.Theanterioronethirdofthenuchalis
asymmetricrelativetotherightside.Thisasymmetryistheresult absentandthelateraledgeoftheleftandrightperipherals1and2
of an anomalous contact between right costal 7, neural 8, and arealsodamaged.Alsomissingfromtheleftsideisthedistalhalf
suprapygal 1. This irregular arrangement was also noted in a ofperipheral6andallofperipherals7–11.Missingfromposterior
specimenofTestudohermanniboettgerifiguredbyLapparentde rightisthedistalhalfofperipheral8,mostofperipheral9,andall
Broin(2006b,fig.10).Thealignmentofneural7withneural8is of peripherals 10 and 11. The pygal bone, suprapygal 2, and the
alsoskewed,butwhetherthisoccurredduringlifeoristheresult lower twothirds of suprapygal 1 are also absent.The rear of the
of preservation cannot be determined. The plastron is nearly turtle has been slightly overthrust onto the fore at the junction
complete, missing only the right lobe of the anal notch, a small between vertebral scute 3 and 4. The plastron is intact but
portion of the left lobe of the anal notch, and the anterior-most severelydeformedandpoorlyprepared.Itwasdeformedthrough
regions of the epiplastra. folding into the visceral space, whereas the xiphiplastron was
FFSM 3446.2.—Specimen FFSM 3446.2 is a well-preserved slightly twisted to the left such that the prominent anal notch no
steinkern with attached plastron (Fig. 3). No cranial or post- longer aligns with the midline.
cranial elements are present. The locations of most visceral SMNS51467.—Thisfragmentaryspecimenconsistsofapprox-
suturesofthecarapacearewellpreservedinthedorsalview.The imately 50 percent of a plastron (Fig. 7). Ventral and visceral
exceptions are the pygal sutures, which are not apparent. Two viewsareavailableandtheepiplastralexcavationisapparent.The
small pieces of the carapace are present but detached from the typicalred-orangemottlingispresentonboththevisceralandthe
main steinkern (not figured). In ventral view approximately 80 ventral surfaces. Missing are the anterior third of the left
percentoftheplastronispresent,andtheepiplastronisremovable epiplastron and a small piece of the anterior right epiplastron.
fromthemainsteinkern.Missingareanteriorandlateralportions Thelateral and posteriorportions of the left and right hyoplastra
oftherighthyoplastron,lateralandposteriorportionsoftheright are missing, as is most of the right hyoplastron. The left
hypoplastron, and most of the right xiphiplastron including the hypoplastronisapproximately75percentintactwiththeanterior
analnotchregion.Theleftlateralportionsofthehyoplastronand and lateral portions missing. Also absent are the posterior two
hypoplastron are missing, as is most of the right xiphiplastron thirds of the xiphiplastron. The left inguinal notch is intact and
including the anal notch region. exhibits a small inguinal scute (not figured).
FFSM3446.3.—Thisspecimenisashellthatisapproximately SMNS 51469.—This is an approximately 70 percent complete
90 percent intact but exhibits overall poor preservation of the shell with some lateral and anteroposterior deformation (Fig. 8).
sulci and sutures (Fig. 4). Most of the neural bones are not No cranial or post-cranial elements are present. As with most of
discernible, including their contacts with the costals. The the other specimens, the shell bones are marked by mottled red-
characteristic red-orange mottling of the Hohenho¨wen site is orange coloration on both carapace and plastron. The anterior
present on the bones. No cranial or post-cranial elements are region of the shell has been polished, whether by geological
present.Missingfromthecarapacearethedistalportionsofright processes or during preparation is unknown. Missing from the
peripheral 10, all of right peripheral 11, all of the pygal, the carapaceontheleftarethedistalportionsofperipherals9and10,
posterior half of left peripheral 10, all of left peripheral 11, the as well as all of peripheral 11. The pygal and much of the
posteriorquarterofthesuprapygal,andthelowerthirdoftheleft suprapygal are absent, as are right peripheral 9–11. The distal
costal 8. The carapace is also missing a small portion of the third of right peripherals 1 and 2 are gone, as are the termini of
anteriormarginofthenuchalboneandapiecefromthemarginof left peripherals 1 and 2. A large indentation is present at the
peripheral1.Theplastronisintactbutthesurfaceiscoveredwith anteriorborderofthenuchalregionandtheleftanteriorsulcusof
chips and divots, likely the result of poor preparation, which vertebral1extendsallthewaytothemarginoftheshellwithno
disrupt most of its surface. A particularly large divot occurs on evidence of a cervical scute. The right terminus of vertebral 1 is
thelefthypoplastronjustanteriorofthecenter;itoccupiesnearly missing,andanunusualsulcustraversesthenuchalbonenearthe
20 percent of the surface area of this bone. terminus. An unusual arrangement of anterior sutures and what
FFSM 3446.4.—This specimen was held in a box labeled appear to be malformed and/or supernumerary bones in the
3446.5 but had 3446.4 written on the steinkern. It is an nuchalregionsuggeststhatoneormoreoftheossificationcenters
approximately 65 percent complete shell (Fig. 5). No cranial or were damaged, perhaps during early development. The posterior
post-cranial elements are present. The characteristic (for Hohen- 20percentoftheplastronismissing,asisaverysmallregionof
ho¨wen) red-orange mottling of the bone surface is present. The the anterior epiplastron.
dorsalviewshowsthecarapacemissingalloftheleftperipherals, UFGC 9.—This is an approximately 70 percent complete
952 JOURNAL OF PALEONTOLOGY, V. 88, NO. 5, 2014
FIGURE 2—FFSM3446.1,Testudoantiqua,middleMioceneofHohenho¨wen,Germany.1–4,photographsandillustrationsofshellin dorsal, ventral,left
lateral,andrightlateralviews,respectively.Notethatwhilethesuprapygalregionisdamaged,thesuturebetweensuprapygal1and2isreadilyapparentinthe
sub-surfacebonethatremains.Heavylinesrepresentsulci,lightlinesbonesutures.Bones:ep epiplastron,en entoplastron,ho hyoplastron,hp hypoplastron,
xi xiphiplastron,nu nuchal,n1-7 neuralbones,c1-8 costalbones,sp suprapygal;scutes:G¼u gular,Hu hu¼meral,Pe pector¼al,Ab abdomina¼l,Fe femoral,
an¼dAn anal,Ce ce¼rvical,P1-4 p¼leuralscutes,M1-1¼1 marginalscute¼s.Scalebar 2cm. ¼ ¼ ¼ ¼ ¼
¼ ¼ ¼ ¼ ¼
CORSINI ET AL.—TESTUDO ANTIQUA FROM THE MIOCENE OF HOHENHO¨WEN 953
FIGURE3—FFSM3446.2,Testudoantiqua,middleMioceneofHohenho¨wen,Germany.1,2,photographsandillustrationsofsteinkernandpartialplastronin
dorsalandventralviews,respectively;3,epiplastroninvisceralview.Heavylinesrepresentsulci,lightlinesbonesutures.SeeFigure2forlabeledbonesand
scutes.Scalebar 2cm.
¼
specimen consisting of a shell with no cranial or post cranial anterior margins of the nuchal, and lower portions of right
elements (Fig. 9). Some deformation is evident in ventral view, peripheral 1 and peripheral 2 are partially degraded. The right
primarily an inward folding of the plastron at the midline and lateral edges of neural 7 and 8 are deformed, and most of right
slight rightward deformation of the carapace. The characteristic costal 8 and the upper half of right costal 7 are damaged. The
red-orange mottling is not immediately evident because a dark plastron is intact except for the anterior margins of the
epoxycoatingcoversmostofthebone;however,inplacesdevoid epiplastron,whichhavebeenslightlydegraded.Theanteriorhalf
of the epoxy, the characteristic red-orange mottling of the oftheleftepiplastronhasalsobeendeformed,beingbenttoward
Hohenho¨wen turtles is apparent. Missing from the carapace are the carapace. The xiphiplastral lobes are largely intact but have
thenuchal,alloftherightperipheralsexcerptpartofperipheral8, sufferedsomeminordegradationatthemargins.Thewidespace
the right half of the pygal, and right peripherals 1, 2, 8, and 9. betweentheterminusofthepygalandtheposteriormarginofthe
Mostofthedistalportionsofleftcostal1andrightcostals1–7are xiphiplastron (i.e., the relatively short posterior lobe of the
missing,asaretheanteriortwothirdsofneural1andtheanterior plastron) suggests that this turtle is a female.
portionsofperipheral3.Afragmentcontainingthelefthalfofthe Overalldescription.—Testudoantiquaisamediumsizedturtle.
pygal,peripheral11,andperipheral10isdetachedfromthemain We were unable to identify any unambiguous autapomorphies
carapace but held in place by the matrix (not figured). The that characterize the species. Those specimens sufficiently
plastron is missing nearly all of both epiplastra except a small complete for measuring range between 18–21 cm in length and
posterior piece of the left bone. Anterior and lateral sections of 14–17cminwidth.Sixspecimens(FFSM3446.1,FFSM3446.3,
therighthyoplastronaregone,asarelateralportionsoftheright FFSM 3446.4, MT PAL.2012.0.10, UFGC 9, and SMNS 51469)
hypoplastron. During preservation, the xiphiplastron and part of are sufficiently preserved to show that T. antiqua had a high-
theleftposteriorhypoplastrondetachedandmigratedinwardand domed aspect with height to width ratios of between 0.59 and
posteriorly while remaining fixed within the matrix. 0.74.Sexisdifficulttodetermineformostspecimens,becauseof
MT PAL.2012.0.10.—The neotype, this well-preserved speci- damage and deformation, but at least two of them (UFGC 9 and
menisrepresentedbyanearlyintactshell(Fig.10).Thecarapace MT PAL.2012.0.10) appear to be female based upon the
is deformed laterally toward the right, and is missing on the left relatively long distance between the pygal and the posterior
peripherals9,10,mostof11,andthedistalthirdofperipheral8. margin of the plastron.
The central parts of the suprapygal region are cracked and Carapace.—Inthosespecimenswithoutextensivedeformation
chipped. The pygal is present, but significantly deformed. The ordegradation (FFSM 3446.1,FFSM3446.3, SMNS 51469,MT
954 JOURNAL OF PALEONTOLOGY, V. 88, NO. 5, 2014
FIGURE4—FFSM3446.3,Testudoantiqua,middleMioceneofHohenho¨wen,Germany.1–4,photographsandillustrationsofshellindorsal,ventral,right
lateral,andleftlateralviews,respectively.Heavylinesrepresentsulci,lightlinesbonesutures.SeeFigure2forlabeledbonesandscutes.Scalebar 2cm.
¼
CORSINI ET AL.—TESTUDO ANTIQUA FROM THE MIOCENE OF HOHENHO¨WEN 955
FIGURE5—FFSM3446.4,Testudoantiqua,middleMioceneofHohenho¨wen,Germany.1,2,photographsandillustrationsofshellindorsalandventralviews,
respectively.Heavylinesrepresentsulci,lightlinesbonesutures.SeeFigure2forlabeledbonesandscutes.Scalebar 2cm.
¼
PAL.2012.0.10) it is apparent that the carapace broadens other such that the anterior and posterior borders of the cervical
posteriorly with the widest point occurring at the apices of arenearlythesamewidth.Thecervicalscutesoftheseriesrange
inguinal notches. In two of these specimens, FFSM 3446.1 and between 25–35 percent length of the nuchal bone. The neural
FFSM 3446.4, the anterior border of the carapace is virtually countcanbedeterminedineightoftheninespecimens;itvaries,
undamaged, showing it to be slightly incut beginning at a small withsixofthespecimensexhibitingeightneurals(FFSM3446.1,
protrusionwheretheposteriorsulcusofmarginal1intersectsthe FFSM 3446.4, SMNS 4450, MT PAL 2012.0.10, UFGC 9) and
border of the shell. The cervical scute is well developed, longer two of them exhibiting seven neurals (FFSM 3446.2, SMNS
thanwide(intact inFFSM3446.1,FFSM3446.3,FFSM3446.4, 51469). Neural 1 is always tetragonal and lozenge-shaped
MT PAL.2012.0.10), and the lateral sulci are parallel to each (roughly square or rectangular) with anterior and posterior
956 JOURNAL OF PALEONTOLOGY, V. 88, NO. 5, 2014
FIGURE 6—SMNS4450,Testudoantiqua,middleMiocene ofHohenho¨wen, Germany. 1–4, photographs and illustrations ofshellin dorsal, ventral,right
lateral,andanteriorviews,respectively.Theanteriorviewshowsthethicknessoftheepiplastrallip.Heavylinesrepresentsulci,lightlinesbonesutures.See
Figure2forlabeledbonesandscutes.Scalebar 2cm.
¼
CORSINI ET AL.—TESTUDO ANTIQUA FROM THE MIOCENE OF HOHENHO¨WEN 957
FIGURE7—SMNS51467,Testudoantiqua,middleMioceneofHohenho¨wen,Germany.1–4,apartialplastroninventralview,ventralillustration,visceral
view,andanteriorview,respectively.Heavylinesrepresentsulci,lightlinesbonesutures.SeeFigure2forlabeledbonesandscutes.Scalebar 2cm.
¼
borders of equal, or nearly equal width. Neural 2 is always specimens slightly narrower anteriorly. This scute is roughly
octagonal,whereasneural3isalwaystetragonal.Thesubsequent hexagonal, though in some cases scute is narrowed laterally and
neurals vary greatly in shape amongst the population, being deviates from a symmetrical hexagon. Vertebral scute 3 is
usuallytetragonal,hexagonal,oroctagonal.Theposteriorneurals hexagonalwithequalanteriorandposteriorspans.Vertebralscute
oftwoofthespecimens(FFSM3446.3,SMNS51467)cannotbe 4 is narrower posteriorly than anteriorly (intact in all except
distinguished. The neural formulae are as follows: FFSM3446.1 FFSM3446.2),althoughthedegreeofnarrowingvariesfrom61–
(4/8/4/8/4/6/?/?), FFSM 3446.2 (4/8/4/8/4/8/4/?), FFSM 3446.3 80percent(posteriortoanteriorborderratio).Vertebralscute5is
(4/?/?/?/?/?/?/6), FFSM3446.4(4/8/4/8/4/6/6/6),SMNS 4450 (4/ much narrower anteriorly than posteriorly, and the lateral sulci
8/4/8/4/6/6/6), SMNS 51467 (?/?/?/?/?/?/?/?), SMNS 51469 (4/8/ descendthroughtheeighthcostaltocontactthepleuralmarginal
4/8/4/8/6/-), UFGC 9 (?/8/4/6/6/6/6/6), MT PAL.2012.0.10 (4/8/ sulcus in all cases except FFSM 3446.3 where the right sulcus
4/8/4/6/6/6).Vertebralscute1ispresentinfiveofthespecimens, crosses into costal 7 prior to contact with the pleural marginal
andinallcasesshowsthecharacteristic‘oniondome’shapethat sulcus. Pleural 1 always contacts marginal scutes 1–5, and
occursinallTestudosp.andIndotestudosp.anddoesnotcontact slightlyoverlapsthenuchal.Thereareeightcostalsandcostals2–
marginalscute2.Insomecasesthisscuteislongerthanwide,and 6 are moderately wedged (alternating converging and diverging
in others it is as long as it is wide. Vertebral scute 2 is in all fromthemidlinetothepleuralmarginalsulcus).Fourofthenine