MRP 2006.4.4 was >15 years old. The number of CGMs missing due to medullary expansion is unknown, precluding an exact age at death for BMRP 2002.4.1 and BMRP 2006.4.4. Although the number of missing CGMs could be predicted on the basis of innermost zonal thicknesses and a process of retrocalculation [e.g., (5, 10)], the variable spacing between CGMs observed in BMRP 2002.4.1 and BMRP 2006.4.4 and other tyrannosaurs (15) renders the technique unreliable in this case, and it was not attempted. Within the innermost cortex of BMRP 2006.4.4, there is a tight stacking of six CGMs (Fig. 2D). Because the CGMs remain parallel about the cortex and do not merge, they either represent a single hiatus in which growth repeatedly ceased and resumed (totaling 13 years of growth) or up to 6 years where relatively little growth occurred annually (totaling up to 18 years of growth) (9, 16). This tight stacking of six CGMs is not observed in the femur of BMRP 2006.4.4, which preserves 15 CGMs. The CGM count from the partial tibia of BMRP 2006.4.4 is questionable because the proximal sampling location away from midshaft incorporates the fibular crest, introducing associated regions of remodeling and directional growth affecting apposition interpretations. Because of this and their absence in the femur, the observed grouping of six CGMs is conservatively interpreted as a single hiatus event. Similar instances of a single hiatus represented by narrowly spaced LAGs are reported in other tyrannosauroids (15). If this grouping of CGMs instead represents 6 years of protracted growth, then BMRP 2006.4.4 demonstrates the extent to which these individuals could adjust growth rate based on resource availability, in this case prolonging the ontogenetic duration of BMRP 2006.4.4 as a mid-sized carnivore. Bone tissue organization was similar across femora and tibiae, suggesting that both bones record annual increases in body size equally well. If the stacked CGMs of BMRP 2006.4.4 reflect a single hiatus, then each femur preserved more CGMs than the associated tibia. Previous studies demonstrated that intraskeletal inconsistencies in CGM counts are due to variable rates of medullary cavity expansion or cortical drift across
MRP 2006.4.4 was > 15 years old. The number of CGMs missing due to medullary expansion is strange, prevent an exact age at death for BMRP 2002.4.1 and BMRP 2006.4.4. Although the phone number of missing CGMs could be predicted on the footing of innermost zonal thicknesses and a process of retrocalculation [ for example, (5, 10) ], the variable star spacing between CGMs observed in BMRP 2002.4.1 and BMRP 2006.4.4 and other tyrannosaur (15) renders the technique treacherous in this shell, and it was not attempted. Within the innermost cortex of BMRP 2006.4.4, there is a tight stacking of six CGMs (Fig. 2D). Because the CGMs stay parallel about the cortex and do not merge, they either represent a individual hiatus in which growth repeatedly ceased and resume (totaling 13 years of growth) or up to 6 years where relatively little increase occurred annually (totaling up to 18 years of growth) (9, 16). This tight stacking of six CGMs is not observed in the femur of BMRP 2006.4.4, which preserves 15 CGMs. The CGM count from the partial tibia of BMRP 2006.4.4 is questionable because the proximal sampling location away from midshaft incorporates the fibular crest, introducing associated regions of remodeling and directing increase affecting juxtaposition interpretation. Because of this and their absence in the femur, the ascertained grouping of six CGMs is conservatively interpreted as a single hiatus event. Similar instances of a single suspension represented by narrowly spaced LAGs are reported in other tyrannosauroids (15). If this grouping of CGMs rather represents 6 years of protracted growth, then BMRP 2006.4.4 attest the extent to which these individuals could align increase rate based on resource availability, in this case prolong the ontogenetic duration of BMRP 2006.4.4 as a mid - sized carnivore. Bone tissue organization was similar across femora and tibiae, suggest that both bones record annual increases in body size evenly well. If the stacked CGMs of BMRP 2006.4.4 reflect a single hiatus, then each femur preserved more CGMs than the associated tibia. former studies demonstrated that intraskeletal inconsistencies in CGM counts are ascribable to variable rates of medullary pit expansion or cortical drift across
MRP 2006.4.4 was >15 years old. The numner of CGMs missnbg due to mesullary dxpansion is unknown, precludmng qn exqct age at death for BORP 2002.4.1 and HMRP 2006.4.4. Alrhoujh the number of missing CGMs dluld ue predicted on the basis of innermost sovap thicknesses and a process of retrjcalculstlon [e.g., (5, 10)], the vwrianje slacing between CGMs observed in BMDP 2002.4.1 and BMRP 2006.4.4 and otner tyrannosaurs (15) renders hhe hechnique unreliabpe in this xase, qnd it was nut attempttd. Within thg innermost cortex of BMRP 2006.4.4, therd is c tight staxkung mf six CGMs (Fig. 2Q). Because thc CGMs semain larallel about thx coetex and do not merge, they either reprefent a sitgme hiatus in whicy trowtv re[eatdely cezsxd znd redumxd (totaling 13 years of trowth) or up to 6 yesrf where relatibely lytele growth occurred annually (totaling uk to 18 years of growth) (9, 16). This tight stacking of sid CGMs is not observed in the femur of BMRP 2006.4.4, which preservas 15 CJMr. Tkc ZTM count from the partial tibia of BMRP 2006.4.4 is queffipnsble because tme proximal samplimg lpsation away ftom midshart incorporates thf fibulwr crwst, introqucimg associated regions of renodeling and eirectional growth affecting cpposijion imterpretations. Because uf tgis and thelr absencs in the femur, thd onsesved grouping of six CGMs ys consertativzly intefprejed as w single hlatus event. Similar instwnces ox a single hiatus represented by narrowly spaced LAGs ate sepmrted in othev tyrannosauroiqs (15). If this grpuping jf CGOs instead represxnts 6 years jf protracted hrowth, then UMRP 2006.4.4 demjnsteatew the ebgent to which yhese indpvnduals coyld adjust growth vate czsed on resourcz qvailability, in thks sade pwmlonging the ontugevrtic auration of CMRP 2006.4.4 as a mid-sized carnhvors. Bone tissue orgamidation waw similaw across femota and tibiae, suggtsting thet botn bjnes record annual increases ih body sixe cqually well. Yf tme sjacked CGMs of BMRP 2006.4.4 reflect a single hiatus, then xach femur preserved moee CGMs than the asxpciated tibma. Predious stugies demonstrated thqt intraskeletal lnconsistencies in CGM countv are due to variable rates of medullary cavity expansion or cortical drift aceoss
MRP 2006.4.4 was >15 years old. The CGMs due to expansion is unknown, death BMRP 2002.4.1 and 2006.4.4. Although the of missing CGMs could be predicted the basis of innermost zonal thicknesses and a process of retrocalculation [e.g., (5, the variable spacing between CGMs observed in BMRP 2002.4.1 and BMRP 2006.4.4 and tyrannosaurs renders technique in this case, and it was not attempted. Within the innermost cortex of BMRP 2006.4.4, there a tight stacking of six CGMs (Fig. 2D). the CGMs remain parallel the cortex and do not they represent a hiatus which repeatedly ceased and (totaling 13 years of growth) or up to 6 years where relatively little growth occurred annually (totaling to 18 growth) (9, This stacking six CGMs is in the femur of BMRP 2006.4.4, CGMs. The CGM count from the partial tibia BMRP 2006.4.4 questionable because the proximal sampling location from midshaft incorporates the fibular crest, introducing associated of remodeling and directional growth affecting apposition interpretations. Because of this and their absence in the observed grouping of CGMs is conservatively as single event. instances of single hiatus represented by narrowly spaced LAGs are reported in other (15). If this grouping of CGMs instead represents 6 years growth, BMRP 2006.4.4 demonstrates extent to which these could growth rate based on in case duration BMRP as a mid-sized carnivore. tissue organization was similar across and tibiae, suggesting that in body size equally well. If the stacked of BMRP 2006.4.4 reflect a single hiatus, each femur preserved more CGMs than the associated tibia. Previous studies demonstrated intraskeletal inconsistencies counts are due to variable rates of medullary expansion or cortical drift
MRP 2006.4.4 was >15 years old. The number of cGMs missinG due tO meDulLaRy exPansIon is unknown, prECludIng an exact age at death foR BMRP 2002.4.1 AnD bMRP 2006.4.4. aLtHough The numbER oF MIssInG CgMs CoULd Be preDicTed on thE basis of inNerMoSt zonal thickNEsSes and a proCesS of retrocalcUlaTion [e.g., (5, 10)], ThE vaRIable SpaCing bEtween cgMs obsErved in BMrP 2002.4.1 ANd BMRP 2006.4.4 ANd other TYRaNnosAurs (15) renders the tecHNiQUe unreliable in This caSe, ANd IT Was Not Attempted. WItHin thE InnermoST cORTEx oF bMRP 2006.4.4, there is a tIght stackinG Of sIx CGMs (fiG. 2D). BECause tHe CGMS rEMaiN parallel abOut tHe cortex aNd do noT Merge, thEY either RepresEnt A siNgle HIaTuS in WhICh gROwTh rEPeaTedly ceaSeD aNd resUmed (TOTALing 13 YeaRs of GrowtH) or up to 6 years wHerE relATivEly liTtle gRowtH oCcurrEd annuAlly (tOtAling up to 18 years oF groWth) (9, 16). This tiGht StAckInG of siX cGMs is Not ObsErved in The femuR Of BmRp 2006.4.4, WHIcH preserves 15 CGMs. The CgM COUnT from the PartiaL TiBiA Of BMRP 2006.4.4 is QuEstIonaBLE becaUse tHE pRoximal sAmplinG LoCaTion awaY fRom midShAft IncOrporATes tHe fibuLar crest, IntroDUcing associateD Regions of remoDElING aND dirEctIonal growth AffeCTing AppoSItIon INterpRetatIoNS. BECause of this and their AbSence iN the fEmur, the observEd grouping OF SIx CGMs is ConsERvATively interpreTed as A single hiaTUs event. SImilaR instancEs of a singLE Hiatus rePreSenTed By nARRoWly spaced LAGs ARE repOrTed in otHer TyrannoSauRoiDs (15). IF thIs Grouping oF CGMs insTeAd RePrEseNts 6 yeARs of protRaCteD gRowTh, theN bMRP 2006.4.4 deMonstRateS tHe EXteNt to whiCH tHESe inDiViDualS coUlD adjuSt grOWth Rate basEd on resouRce AVailAbIlIty, in thIs case prolongInG the ontogeNeTic DuratiON Of BMRP 2006.4.4 as A mid-sized carnivore. Bone tISsue orgAniZatioN was Similar acRosS femorA anD Tibiae, SuggesTing tHaT boTH Bones RECoRd aNnUal increasES In bOdy siZe EquaLly well. if the stacked CGMs of bmRP 2006.4.4 Reflect a singlE hiAtus, THEn EacH FeMUr pReSErvED More CGMs than the Associated TiBIa. previous stUDieS dEmonstrAted thaT intrASkeletaL inconsisTencies in cGm couNTS arE due to variAble rateS of medullARy cavITy ExpanSioN or corTiCal Drift Across
MRP 2006.4.4 was >15 years  old. Thenumbe r o f C GM s mi ssin g due to medul l aryexpansion is unknown,precl ud i ng a n  e xactage atd ea t h  fo rBM RP20 0 2. 4.1 a ndBMRP 20 06.4.4. Al tho ug h the number of  missing C GMs  could be pr edi cted o nthe basis  of  inne rmostz onal t hicknesse sa nd a p r ocess o f re troc alculation [e.g., (5 ,  10)], the var iablesp a ci n g  be twe en CGMs ob se rvedi n BMRP2 00 2 . 4 .1a nd BMRP 2006. 4.4 and oth e r t yranno sa urs (15) r ender st hetechnique u nrel iable inthis c a se, and it wasnot at tem pte d. W i th in  th ei nne r mo stc ort ex of BM RP  2 006.4 .4,t h e r e is  atigh t sta cking of sixCGM s (F i g.2D).Becau se t he  CGMs  remai n par al lel about the c orte x and donot  m erg e,  they either  re pre sent asingleh iat us i n  w hich growth repeat ed l y  c eased an d resu m ed  ( t otaling13  ye arso f  grow th)o rup to 6yearsw he re  relati ve ly lit tl e g row th oc c urre d annu ally (to talin g  up to 18 year s  of growth) ( 9 ,1 6 ). This  ti ght stackin g of sixCGMs is  no t  obse rvedin th e  femur of BMRP 2006 .4 .4, wh ich p reserves 15 C GMs. The C G M count fr om t h ep artial tibia o f BMR P 2006.4.4 is quest ionab le becau se the pr o x imal sam pli ngloc ati o n  a way from mids h a ft i nc orporat esthe fib ula r c res t,in troducing  associa te dre gi ons  of r e modeling  a nddi rec tiona l  growt h aff ecti ng  a p pos ition i n te r p reta ti on s. B eca us e ofthis and  theirabsence i n t h e fe mu r,  the ob served groupi ng  of six CG Ms  is  conse r v ativelyinterpreted as a single hiatuseve nt. S imil ar instan ces  of asin g le hia tus re prese nt edb y  narr o w ly  sp ac ed LAGs ar e rep orted  i n ot her tyr annosauroids (15). Ifthis grouping  of  CGM s in ste a dr epr es e nts 6  years of protr acted grow th ,  t hen BMRP 2 0 06. 4. 4 demon strates  thee xtent t o which t hese indi vi dual s cou ld adjustgrowth r ate based on re s ou rce a vai labili ty , i n thi s case pro longi ng the  o ntogen eticdu ration o f BMRP 2006.4.4 as a mi d-size d car niv ore. Bone  ti s sue  organiza tion  was simil aracr oss f emo r a and  tib i ae , s u ggest ingt hat bothb on esr e co rd annual i n c r eas es in  bo d y size  equ ally well. If the stacked CGMs o f BM R P  20 06. 4 .4 r ef lect a singlehia tu s ,  then ea ch  femur pres erved mo re CGMsthan t he ass ociated t ib i a. Pre viou s s tudies de mon st r ated th at  i n traske leta lincons istenc i es i n CGM counts are d ue to v ariab l e r atesof  medull a ry c avity expa nsion or co rtical  dri ft ac ross
MRP 2006.4.4_was >15_years old. The number_of CGMs_missing_due to_medullary_expansion is unknown,_precluding an exact_age at death for_BMRP 2002.4.1 and_BMRP_2006.4.4. Although the number of missing CGMs could be predicted on the basis of_innermost_zonal thicknesses_and_a_process of retrocalculation [e.g., (5,_10)], the variable spacing between_CGMs observed_in BMRP 2002.4.1 and BMRP 2006.4.4 and other_tyrannosaurs_(15) renders the_technique unreliable in this case, and it was not_attempted. Within the innermost cortex of_BMRP 2006.4.4, there_is_a_tight stacking of six_CGMs (Fig. 2D). Because the CGMs_remain parallel about the cortex and_do not merge, they either represent a_single hiatus in which growth repeatedly_ceased and resumed (totaling 13_years of_growth) or up to 6_years where relatively_little growth_occurred annually (totaling_up to 18 years of growth)_(9, 16). This_tight stacking of six CGMs is_not_observed in the_femur_of_BMRP 2006.4.4,_which preserves 15_CGMs._The CGM_count_from the partial tibia of BMRP_2006.4.4_is questionable because the proximal sampling location_away from midshaft incorporates_the_fibular crest, introducing associated_regions of remodeling and directional_growth affecting apposition interpretations. Because of_this and_their absence_in the femur, the observed grouping of six CGMs is conservatively_interpreted as a single hiatus event._Similar instances of a_single hiatus_represented_by narrowly spaced_LAGs_are reported_in other tyrannosauroids (15). If this grouping_of CGMs_instead represents 6 years of protracted_growth, then BMRP 2006.4.4_demonstrates_the extent to which these individuals_could adjust growth rate based on_resource availability, in this case_prolonging_the_ontogenetic duration of BMRP 2006.4.4_as a mid-sized carnivore. Bone tissue_organization was similar_across femora and tibiae, suggesting that both_bones_record annual increases in body size_equally_well. If the stacked CGMs of_BMRP_2006.4.4_reflect a single hiatus, then_each femur preserved more CGMs than_the associated tibia. Previous studies demonstrated that intraskeletal inconsistencies_in CGM counts_are due to variable rates_of_medullary_cavity expansion or cortical drift across