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A Fossil Trionychid Turtle F Rom the Early Tertiary Chuckanut Formation

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A Fossil Trionychid Turtle F Rom the Early Tertiary Chuckanut Formation GeorgeE- Mustoe and SamuelP. Girouard,Jr., Geo ogy DepartrnentWestern Wash ngton Universty, Be ngharn Washngton 98225 A FossilTrionychid Turtle f rom the earlyTertiary Chuckanut Formation of NorthwesternWashington Abstract \Vrshington s \'e(cbrute fossilfecordi\ so sparsethate!en, discover] desenescarcful scnrtiny.AfossilizcdturlecaraDace tbund in 1960in fluviatilcsard stone of theChLrckanut Formaiion of wcnem WhatcomCounly $ a, ,,fig ini | 1idtnritrril.6aillcmheru1 the Tesludinoideasupedrmil). and an adjrcent bonefragmcnt was consideredtobceridence ofsome largeranimal. Rc cianina- tlon indicalcs (har the cafapacercpreserls a member of thc Trionychidae (soft shclled tufiles). Compulerired axial tomogmphv \Lrggeststhal the neafb) bone probably came from $c tu e. Trionychid fossils are abrndant in carl] Tefiiary rocks. bul rheir remarnsIa!c nol beenfound in younger rocks in ihe north$en resion. The only cxranrtrionychid found west of rhe MississipDi is ./l'i.,ir r \prlrl,rJ'ur.which inhabitssmall areasof eastemMontana and the Colorado Rivef fe8inn ot rh( "ourh$e\r. t'hr\rurrrrr- llon in geographicrange probablt resultedli|)lr global cooling lhar surted in the latc Eocene,and by habiut destructioncauscd b,"-episodes of nrounlainbuilding ihat begana lc$ million yearslarcr. Introduction raphy ofthe PaddenMenlber. but its age is prob ably lateEocene based on stmcturalrelationships The faunalrecord from the earlyTcftiary is sparse andpaleoclimatic evidence. Mustoe and Pevear evcnthough extensive sedimentary deposits liom (1983)described the poorly-preservedcarapace thisperiod are exposed on bothsides of theCas- asa memberof the SuperfamilyTestudinoidea, a cadeRange. These beds of shale,siltstone. sand- large group that includes pond turtles and land stone,conglomerate, and coal were depositedby tortoises.They guessedthat bone fragment ex- meanderingd!e$ that flowed westward across posedin cross-sectionview on one side of the thc broad lloodplain that coveredmuch of west- specimenwas part ofa rib bonefrom somelarger em and centralWashington prior to the uplift of animal. Becauseturtles havc ribs that are fused the North Cascades.Of thesedeposirs, the 6,000 to the adjacent bony carapacelayer. this bonewas nr thick ChuckanutFormation has reccived the presumedto havg come either fiom bonesof a lrroslattention trom geoscientists(Pabst 1968; land-dwellingmammal that werewashed into the Johnson 198.1;Musloc 1993; Musroe and fluvial sediments.or to be skeletalrernains front Gannaway 1997).Plant remainsare abundantin an aquatic reptile. The latter possibiliry is con Chuckanutstrata and animalsare representedby sistentwith discoveriesoffossil crocodilesin the a varictvoftracks. The only vedebratebody los- GreenRiver Forrration of Wyoming (Grande sils that ha\e been fbund are the impressionof 198,+),correlative in age to the ChuckanutFor- the interior surfaceof the carapaceof a tu le and nlation. Becausethe fossil was softer than the r nearbybone fragrnent (Figures l.2). enclosingmatrix, the decision was made not to The l8 x l3 x 7 cm spccimenwas collected in attempt to reveal the bone by mechanicalexca 1960during constructionexcavations south of vationfor fearof destroyingthe specimen.In 1996 BelJinghamat Clark Point(T 37 N. R 2 E. SW geologystudent S. P Girouard.Jr, re-eramined comcr Sec 13)but recognitionof the fossilas a the specimenand decided that both of Mustoe turtle campacedid not occur until two declrdes rntl Perear'.tl98.lt interpretction\uere incor- later when a visiting geologist noticed the rock rect. This paperpresents his findings. at the Clark family home (Guthrie1981). Bed- Anatomica Features rock at Clark Point consistsof arkosicsandstone of the Carapace ol the PaddenStratigmphic Menber, the young- Althoughtheimpression ofthe carapacepresen es estsubunit ofthe ChuckanutFormation. Johnson relativelytew anatomicaldetails, sevcrll char ( 1984)and Mustoeaod Gannaway(1997) pre acteristicssuggest that the spccimenreprescnts sentcdconllicting interpretationsfor the stratig- a mcmber of the Superfamily Trionychidae NorthwestScience. Vol.75, No.3,2001 2ll O llr0l br_rtrc Nodhsc{ SoenrifirAsociarnri Allnghr.eserl.d Figure L Chuckanut tuftle fossil. showlng inrprcssionol lenlral surfaceof carapace.Broad corrugalionsrepre- scnt iusing of thc ribs to iblm a conrinuousbon) la,ver.These corrugalions meet :rt a midline axis rhat represeDtserodcd ncural archesoi lhe lerlebfal column. Scalebar = ! cn. Photosused fbr figs. I & 2 ha\'e beenreversed from rjght to lcli 1()a]lo$ easiercomparison $,ith the radiographin lig. 6. Figure2. Srde!ie\i ofspecinren.with bonefragnren! arkcdbvinowlsamescaleasFigurel). 212 Mustoeand Girouard Figure 3. Iniernal!ie\| oft\\'o t)pes of carapacearchitecrurc. Lcflr T.a.renr5 r.rrr1rr. showingthe dng ofperipheral platesthal sunoundthe uppef shellin rll runles ercept liony chids.Righti /-ir-re,r_!r/,rddra . aoion,vchid.Adapted liom Me)lan d9li7). laginousnrtrgin typicallycomprises approximately one half of the total carapaceleDgth. In many speciesthe distal endsof the ribs project beyond the perimeterof the central bony disk, but other trionychidslack theseprojections (Meylan, 1987). An importantdistinguishing characteristic of trionychidsis thc absenceol epidermalplates (scutes).Instead, the shell'sexternal surtace is covered with leathery skin (Figure 5). This ab- senceof homy scutesis sharedonly by the huge leatherbacksea tunlcs, Family Demochelyidae (Emstet al. 1994). Figure,1. Fossil soli shclledturtle 7'il)lirf ,r? rrtlnrnrs fron Eoccnc shalcat N{essel.German}. sho\\ing dorsal !iew ofcarapacc.Thc bump! lerrure ofthe dorsal su acehelps thebon) plalesprovide astrong s b- st|:lleibr the overlling ]calhcry carapace.as illus- In,rrJ In Fi;Lr< 5. Th( (-hu.l'enur.nc\ -en. f pears quite dillerenl liom cilhcr of these figures becluse the lbssil sholvsrhc \ cnlral suriace.Photo courlcsy oi ScnckcnbcrgMuseum. Frankfurt. (sofi-shelledtuftlcs). Other turtles have carapaces (Easrem thalhave dgid margirs composedof20 to 22 bony Figurc 5. The livnrg lrbr]I spi/rildrr Spiny Soit shell) sho$,sthe leathef) carapacesudace charac' plates.but triorychjds have flexible margins and teristic of membersof the Trionichidae.Photo b) lack peripheralbones (Figures 3,4). The carti' Fl .1.Obsr. Trionychid Tu le from Tefiiary ChuckanutFormation 213 Possibly the Chuckanut fbssil representsan among trionychidscarapace shape remains rela incompletecarapace where periphelal bones were tively constantduring growrh. cither disarticulatedprior to fossilization,or de- stroyedlater by erosion.A ntore likely interpre- Ev dence f rorn Computerized Axia talionis that the absence ofthese marginal plates Tomography is evidencc that the fossil is from a trionychid. Thc sandstoneblock also preserles a tbssilbone Be.ru.e thc \f\crimen pre.erre\ rn irnpre.sion visible in crosssection as a flattenedasymmetric lefl by the interior surtaceof the carapace.it is 26 x 6 mm oval, compdsedof a 0.5 mm thick not possibleto determinethe natureof the origi- cofiical layer sur:roundinga porous ccntral zone nal outer surface.However, a partially preserved (Figure2). ln May, 1996a computerizedaxial layer of mineralized bone bears no trace of im- tomography analysis of the specimenwas per- pressions(sulci) that comrnonly fbrm alongseams formed using a GeneralElecrric Model 9800 CT betu'eenadjacent scutes. The shallow convex scanner.The valueofCT technologyfor the study corrugatedexterior sulfacebears a strongresem of vertebratefbssils was first demonstratedby blanceto shell shapesofextant speciesof lrloa_rx Conroy and Vannier (198.1),and the technique and its kin. though some other types of turtles wasquickly adoptedby veflcbratepaleontologists havesinilar contours. (for a detailedoven,iew seeClark and Morrison The l2 cm lengthof the Chuckanutfossil is 1994).The Chuckanutspecimen was subiected smallcompared to carapacclengths ofmostmodem to 18 scanson the coronal (anteriorto posterior) andtbssil trionychids.LiLrge extant members have andsagittal (nredial to lareral)planes, usually at bony disks that excecd50 cm, and for most spe, an inteNal of 3 n]m betweensuccessivc scans and clesthe adultshave carapacial disk lengthsof 20- at an accelentingvoltage of 120KV ar 100-170 30 cm. Of 23 living species,only five havedisks MA anda scanrate of 37.5mm/s. Because ofthe lessthan 20 cm (Meylan 1987).The Chuckanut density of the matrix the scanimages arc some- specimgnmay representan unusuallysmall vari- what indistinct,butrcsolution is sufficientto show ety, but it may also be remains of a juvenile be- the generaishape and dimensionsof the hidden causcin contrastto many other types of turtles. bone fragment (Figure 6). Figure 6. Tracingsm.tde fioln five CT scansshow successivecross sectionvie\\'s ofboDe fragmen! that exrendsapproximately l0 mm irlo lhe sandslonematrix Photo showsactual CT scanimage al 12 nm deprh.with included bone marked by affow. The undulating surfacc al lhe top ofthe siab representsthe carapaceimpfession. The size and flarcned shape suggcstthat the bone is probably either a fragnrcn!oLplastron or intemal skeleton. 21,1 Mustoeand Girouard Figure L Sketchesollrionychid rhell architectlrre.shown in \,ennal \ie\l\.,Adapted iiom \{eylan (1987). Lcll: Trio \rJ(t?r. Right I air.l(Drt r p&r.trd . The preservedponion of the bone is a thin. 'ubtriangularstructure lhul e\lend. lor approri- mately30 mm into the matrix. Although an asym- metricflattengd oval crcss-sectioncan be obseryed at its exposureat the surfaceofthe slab.the bone ha: r compres.edtrapezoidrl crors reetiun
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