Phylogenetic RelationshipsN`k_`e<mfclk`feXip9`fcf^p ROMI L. BURKS $ IPDPMBUF DBMNT UIF NJOE ZFU FYDJUFT UIF TFOTFT $IPDPMBUF BMTP VOJUFT DVMUVSFT %JQEQNCV B NPWJF BCPVU B TNBMM UPXO 'SFODI DIPDP MBUF TIPQ NBEF NJMMJPOT JOUFSOBUJPOBMMZ 4UBSSJOH BDUPSTDPOUSJCVUFEQBSUMZUPUIFGJMNµTTVDDFTTCVUUIF GJMN BMTP ESFX TBMJWBUJOH WJFXFST XPSMEXJEF UP UIF NVMUJQMF BQQMJDBUJPOT PG DIPDPMBUF 8JUI JUT TUJNV MBOU QSPQFSUJFT DIPDPMBUF HFOFSBMMZ NBLFT QFPQMF GFFMHPPE$IPDPMBUFNBZBMTPJOTQJSFDSFBUJWJUZ'PS FYBNQMF FWFSZ EBZ CJPMPHJTUT DBO VTF DIPDPMBUFµT GPSUVOBUFQSPQFSUZPGBQQFBSJOHJOEJGGFSFOUTIBQFT TJ[FTBOEDPNQPTJUJPOTUPIFMQTUVEFOUTVOEFSTUBOE CBTJDQSJODJQMFTPGFWPMVUJPO'FXTUVEFOUTIBWFWJTJU FEUIF(BMBQBHPTUPTFF%BSXJOµTGJODIFTCVUOFBSMZ BMM IBWF TBNQMFE B WBSJFUZ PG DIPDPMBUF CBST 6TJOH DIPDPMBUF BT B ²NPEFM PSHBOJTN³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igure 1. Abridged history of the chocolate bar and candies. • 1890 George A. Bayle Jr., of St. Louis, Missouri, packages peanut butter in barrels and sells it for six cents a pound. • 1900 Milton S. Hershey of Lancaster, Pennsylvania, introduces the first HERSHEY’S® Milk Chocolate bar. • 1903 George Washington Carver researches uses for peanuts. Peanut butter introduction to the world occurs a year later at the Universal Exposition in St. Louis. • 1920 Baby Ruth® candy bar is first sold, named for President Grover Cleveland’s daughter. • 1928 Hershey introduced peanut butter cups later named after Harry Burnett (H.B.) Reese, a former dairy employee. • 1930 M & M/Mars introduces the SNICKERS® Bar, named for a favorite horse owned by the Mars family. One candy bar costs a nickel. • 1941 M & M’S® Plain Chocolate Candies (later M & M’S® Milk Chocolate Candies) introduced. INQ UIRY & I N V ES T I GAT I ON <MFCLK@FEF=K?<:?F:FC8K<98I1 8:i\Xk`m\8ggifXZ_kfK\XZ_`e^ • 1954 M & M’S® Peanut Candies introduced. • 1978 REESE’S PIECES candy is introduced. • 1982 REESE’S PIECES candy makes its big screen debut as “E.T.’s” favorite candy in the movie E.T., The Extra Terrestrial. • 1981 A European favorite since 1974, SKITTLES® Bite Size Candies are introduced in the U.S. by M & M/Mars. • 2001 SNICKERS® Cruncher premiers. • 2002 SNICKERS® with Almond Bars replace Mars Bar. • 2004 M & M’S® and chocolate bar merge into M-Azing® Crunchy and Peanut Butter versions. Sources: National Confectioners Association (http://www.candyusa.org/ Classroom/timeline.asp), http://www.peanutbutterlovers.com/history/, M&M Mars Corporation (http://www.mmmars.com/cai/snickers/faq.html#top, http://www.snickers.com/history.asp, and http://us.mms.com/us/about/history/ story/)), and Hershey Foods Corporation (http://www.hersheys.com/consumer/ history.shtml;. All candies are registered trademarks. 41/+ . $74-5 2J& DWTMUT"UQWVJYGUVGTPGFW KU #UUKUVCPV 2TQHGUUQT QH $KQNQI[ CPF .#44; % $1.'5 YCU 8KUKVKPI #UUKUVCPV 2TQHGUUQT QH $KQNQI[ DQVJ CV 5QWVJYGUVGTP 7PKXGTUKV[)GQTIGVQYP6: OFTTFTQFDJBMMZXIFOTIBSJOHTVDIJEFBTXJUIOFXMZNJOUFE VOEFSHSBEVBUFT PS IJHI TDIPPM TUVEFOUT XIP NBZ PS NBZ OPU IBWF FYQFSJFODFE B QSPHSFTTJWF CJPMPHZ DVSSJDVMVN #ZCFF-BOHFO PHYLOGENETIC RELATIONSHIPS 229 Figure 2. Glossary. 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SFTVMUJOHGSPNUIFCSBODIJOHPGGPGOFXUBYBGSPNDPNNPO BODFTUSBMMJOFBHFT $MBTTJGJDBUJPO ° UIF TZTUFNBUJD QSPDFTT PG GPSNJOH BOE PSEFSJOHHSPVQTCBTFEPOTJNJMBSJUZ $POWFSHFODF°TJNJMBSJUZFTQFDJBMMZPGGVODUJPOBQQFBSJOH JOUBYBUIBUEPOPUTIBSFBOJNNFEJBUFDPNNPOBODFTUPS EVFQSJNBSJMZUPTJNJMBSTFMFDUJWFQSFTTVSFT %FSJWFE°BDIBSBDUFSTUBUFUIBUJTBNPEJGJFEWFSTJPOPGUIF BODFTUSBMTUBUF &WPMVUJPO ° DIBOHF JO BMMFMF GSFRVFODJFT JO B QPQVMBUJPO PWFSUJNF &WPMVUJPOBSZTUFQ°BDRVJTJUJPOPSMPTTPGBDIBSBDUFSUIBU DIBOHFTUIFSFTVMUJOHMJOFBHF &YUBOU°MJWJOHUPEBZ 'JUOFTT°NFBTVSFPGPOFµTHFOFUJDDPOUSJCVUJPOUPUIFOFYU HFOFSBUJPOSFMBUJWFUPPUIFSJOEJWJEVBMT )PNPQMBTZ ° TJNJMBSJUZ EVF UP JOEFQFOEFOU FWPMVUJPOBSZ DIBOHF )PNPQMBTZ JT FJUIFS QBSBMMFMJTN JOEFQFOEFOU /FWFSUIFMFTT XF LOPX UIBU ²<O>PUIJOH JO CJPMPHZ NBLFT TFOTFFYDFQUJOUIFMJHIUPGFWPMVUJPO³%PC[IBOTLZ 5IVT UIFOFFEGPSCJPMPHZFEVDBUPSTBDSPTTNVMUJQMFFEVDBUJPOBMMFWFMT UPUBDLMFUIFTVCKFDUPGFWPMVUJPOXJMMPOMZJODSFBTFPWFSUJNF" OFXMZSFOFXFEJOUFSFTUCZTDJFOUJTUTJOUIFXBZUIBUFWPMVUJPOJT UBVHIU"MUFST/FMTPO IBTTQBXOFEBOVNCFSPGIFMQGVM 8FCTJUFTBOEUFDIOPMPHJDBMSFTPVSDFT"*#4 *OBEEJUJPO %BSXJOJBO FYDFSQUT GSPN 6JG 1TKIKP QH 5RGEKGU NBZ IFMQ FMVDJEBUF UIF GPVOEBUJPOBM DPODFQUT PG FWPMVUJPO JF OBUVSBM TFMFDUJPOGJUOFTTBOEBEBQUBUJPO $PTUB )PXFWFSPOMZ BGFXPGUIFTFSFTPVSDFTEFMWFGVSUIFSJOUP²EFTDFOUXJUINPEJGJDB UJPO³%BSXJO BOEJOWFTUJHBUFUIFQSPDFTTPGEFUFSNJOJOH UIF FWPMVUJPOBSZ SFMBUJPOTIJQT CFUXFFO PSHBOJTNT :FU XJUIPVU TVDIVOEFSTUBOEJOHBOJOUSPEVDUPSZFYQMPSBUJPOPGUIF²5SFFPG -JGF³.BEEJTPO XJUIIJHITDIPPMTUVEFOUTPSVOEFSHSBEV BUFTDBOCFBSPVHIKPVSOFZ 230 THE AMERICAN BIOLOGY TEACHER, VOLUME 69, NO. 4, APRIL 2007 HBJO PSSFWFSTBMMPTT /BUVSBMTFMFDUJPO°EJGGFSFOUTVSWJWBMBOESFQSPEVDUJPOPG JOEJWJEVBMTDBSSZJOHBMUFSOBUFJOIFSJUFEDIBSBDUFSJTUJDT .POPQIZMFUJD°BTFUPGUBYBDPOUBJOJOHBDPNNPOBODFT UPSBOEBMMJUTEFTDFOEBOUT /PEF°BCSBODIQPJOUPOBUSFFDMBEPHSBN 0VUHSPVQ ° B UBYPO VTFE UP IFMQ SFTPMWF UIF QPMBSJUZ PG DIBSBDUFST BOE UIBU JT IZQPUIFTJ[FE UP CF MFTT DMPTFMZ SFMBUFEUPFBDIPGUIFUBYBVOEFSDPOTJEFSBUJPOUIBOBOZBSF UPFBDIPUIFS 1BSBQIZMFUJD ° TFU PG TQFDJFT DPOUBJOJOH BO BODFTUSBM TQF DJFTUPHFUIFSXJUITPNFCVUOPUBMMPGJUTEFTDFOEBOUT 1BSTJNPOZ°POFPGTFWFSBMDSJUFSJBUIBUNBZCFPQUJNJ[FE JOCVJMEJOHQIZMPHFOFUJDUSFFT5IFDFOUSBMJEFBPGDMBEJTUJD QBSTJNPOZBOBMZTJTJTUIBUTPNFUSFFTXJMMGJUUIFDIBSBDUFS TUBUF EBUB CFUUFS UIBO PUIFS USFFT 'JU JT NFBTVSFE CZ UIF OVNCFSPGFWPMVUJPOBSZDIBSBDUFSTUBUFDIBOHFTJNQMJFECZ UIFUSFF 1IFOFUJDT°BNFUIPEPGDMBTTJGJDBUJPOCZXIJDITJNJMBSJUZ JTEFUFSNJOFECZDPOTJEFSJOHQIFOFUJDDIBSBDUFSTDIBSBD UFSTSFTQPOTJCMFGPSBOFOUJUZµTBQQFBSBODF $IBSBDUFSTBSF TFMFDUFEXJUIPVUSFHBSEUPBOFOUJUZµTFWPMVUJPOBSZIJTUPSZ 1IFOPUZQJDQMBTUJDJUZ°OPOHFOFUJDWBSJBUJPOJOPSHBOJTNT JOSFTQPOTFUPFOWJSPONFOUBMGBDUPST 1IZMPHFOFUJDUSFF°BEJBHSBNSFQSFTFOUJOHUIFFWPMVUJPO BSZEFWFMPQNFOUPGSFMBUFETQFDJFT 3PPUFE USFF ° B DMBEPHSBN XJUI B IZQPUIFUJDBM BODFTUPS UIBU FRVBUFT UP UIF SPPU UIBU JT UIF OPEF BU UIF CBTF PG UIF USFF 8IFO PVUHSPVQT BSF VTFE UIJT JT UIF OPEF UIBU DPOOFDUTUIFPVUHSPVQTUPUIFJOHSPVQBOEUIVTTQFDJGJFT UIFEJSFDUJPOPGFWPMVUJPOBSZDIBOHFBNPOHUIFDIBSBDUFS TUBUFTDGVOSPPUFEUSFF 4JTUFS HSPVQ ° B DMBEF UIBU JT CFMJFWFE UP CF UIF DMPTFTU HFOFBMPHJDBMSFMBUJWFPGBHJWFOUBYPOFYDMVTJWFPGUIFBODFT USBMTQFDJFTPGCPUIUBYB0SUIFEFTDFOEBOUCSBODIFTGSPN BOPEFPOBDMBEPHSBN 4ZOBQPNPSQIZ°BEFSJWFEPSTQFDJBMJ[FEDIBSBDUFSTIBSFE CZUXPPSNPSFHSPVQTUIBUPSJHJOBUFEJOUIFJSMBTUDPNNPO BODFTUPS 5PQPMPHZ°UIFCSBODIJOHTFRVFODFPGBUSFF 1IZMPHFOZ$MBTTJGJDBUJPO$MBEPHSBNT 5SFFT 8F DPNNPOMZ IFBS UIBU ²B QJDUVSF JT XPSUI B UIPVTBOE XPSET³%BSXJONBZIBWFUIPVHIUUIJTXIFOXSJUJOHIJT1TKIKP QH 5RGEKGU XIFSF IF DIPTF UP JODMVEF POMZ POF EJBHSBN 'SFFNBO )FSSPO %BSXJOµT ²IZQPUIFUJDBM USFF³ JMMVT USBUFE IJT WJFX PG IPX TQFDJFT DIBOHF UISPVHI UJNF 5PEBZ QIZMPHFOFUJD USFFT TFSWF BT WJTVBM SFQSFTFOUBUJPOT PG UIF QPT TJCMF IJTUPSJDBM SFMBUJPOTIJQT CFUXFFO TQFDJFT JF QIZMPHFOZ XIFSF TQFDJFT BU UIF UJQT PS FOET PG USFFT SFQSFTFOU TQFDJFT HFOFSB PS PUIFS UBYB MJWJOH UPEBZ JF FYUBOU PS UIBU NJHIU IBWF HPOF FYUJODU #SBODIFT PG USFFT UIFO SFQSFTFOU BODFTUSBM QPQVMBUJPOTPGTQFDJFTUISPVHIUJNFBOEOPEFTEFTJHOBUFQPJOUT XIFSFPOFTQFDJFTTQMJUTJOUPUXPNPSFEFTDFOEFOUQPQVMBUJPOT 'SFFNBO)FSSPO 3PPUFEUSFFTJEFOUJGZUIFPSJHJOPG UIF MJOFBHF BOE UIF DPNCJOBUJPO PG UJQT CSBODIFT BOE OPEFT JMMVTUSBUF %BSXJOµT DPODFQU PG ²EFTDFOU XJUI NPEJGJDBUJPO³BOEUIFQMBVTJCMFSFMB UJPOTIJQTBNPOHPSHBOJTNT Figure 3. Different ways to classify sets of candies. In the cut-away view: BR = Baby Ruth® RP = REESE’S PIECES HWC = HERSHEY’S® White Chocolate SNC = SNICKERS® Cruncher MMPE = M & M’S® Peanut HC = HERSHEY’S® chocolate bar SN = SNICKERS® HA = HERSHEY’S® with Almonds MMPL = M & M’S® Plain SNA = SNICKERS® Almond Bar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igure 4. Sample phylogenies for sister groups of candy pieces (A) and candy bars (B). Perpendicular lines on branches CF VTFE UP DMBTTJGZ UBYB represent evolutionary steps. SFTFBSDIFSTNVTUDBSFGVMMZ DPOTJEFS PO XIJDI USBJUT UIFZXJMMSFMZ $MBEJTUJDT PS QIZMP HFOFUJD TZTUFNBUJDT BGUFS )FOOJH JT B NFUI PE PG QIZMPHFOFUJD DMBT TJGJDBUJPOUIBUBUUFNQUTUP MJNJU DPNQBSJTPOT XJUIJO HSPVQTUPUIPTFDIBSBDUFST UIPVHIU UP IBWF FWPMWFE XJUIJOBHJWFOHSPVQBMTP LOPXO BT EFSJWFE DIBS BDUFST5IFQSPDFTTPGDMB EJTUJDT XPSLT CZ BOBMZ[ JOH EJGGFSFOU UBYB UP GJOE PCKFDUJWF TJNJMBSJUJFT BOE EJGGFSFODFTCFUXFFOUIFN 0OFUIFOVTFTUIPTFTJNJ MBSJUJFT BOE EJGGFSFODFT UP DSFBUF B USFF TUSVDUVSF JMMVTUSBUJOHUIFNPTUTJNJ MBSUBYB5IFBTTVNQUJPOJT UIBU TJNJMBS UBYB BSF TJNJ MBSCFDBVTFUIFZBSFFWPMV PHYLOGENETIC RELATIONSHIPS 231 TJTUFSUBYBTVDIBTBMMWBSJ FUJFT PG DBOEZ XJUI QJFDFT 'JHVSF " XIJDI BSF BMM EFSJWFE GSPN BO VOLOPXO DPNNPOBODFTUPS Figure 5. Example of MacClade windows and output. Four possible cladograms shown on the left with the most parsimonious tree (13 steps, shown in Tree Window) appearing below the data matrix (seen in Data Editor window). Larger box shows tool palette from MacClade where the arrow is used to move branches to arrive at the tree with the fewest steps. Common ancestor assumed to be a chocolate bar. %BSXJOµTGJODIFTTFSWF BT UIF NPEFM GPS BTLJOH FWPMVUJPOBSZ RVFTUJPOT (SBOU (SBOU CVU MJGF PO UIF (BMBQBHPT NBZ CF UPP GBS SFNPWFE GSPN TUVEFOUT UP NBLF BO JNQBDU4UVEFOUTBOEUFBDI FST HFOFSBMMZ SFBDU RVJUF GBWPSBCMZ UP DIPDPMBUF BOE NBZFWFOLOPXTPNFUIJOH BCPVU JUT IJTUPSZ 'JHVSF XIJDI JT GBTDJOBUJOH BMUIPVHICFZPOEUIFTDPQF PG UIJT QVCMJDBUJPO *O UIF BHF PG BMUFSOBUJWF UFBDIJOH TUSBUFHJFT GPS FYDFMMFOU SFTPVSDFT TFF #FBO .D,FBDIJF .JDIFM UIFMFDUVSFBQQSPBDI GPS DPNQMFY JEFBT TVDI BT QIZMPHFOFUJDT PGUFO MFBWFT TUVEFOUT QJOJOH GPS QSBDUJ DBM BQQMJDBUJPO *O UIF GPM MPXJOH FYFSDJTF TUVEFOUT VTF B HSPVQ PG DBOEZ CBST UPMFBSOBCPVUQIZMPHFOFUJD QSJODJQMFT BOE IPX TDJFO UJTUT XPSL UP VOEFSTUBOE UIF SFMBUJPOTIJQT UIBU FYJTU CFUXFFO PSHBOJTNT 'SBNJOH B EJTDVTTJPO BCPVU FWPMVUJPO CZ VTJOH DIPDPMBUF TJNQMZ QSPWJEFT B CPVOEBSZ GPS TUVEFOUT UP GSBNF UIFJSDPODFQUJPOTBCPVUBUPQJDUIBUJTPGUFOEJGGJDVMUUPBQQSPBDI BOE UFBDI 5IF FYFSDJTF EFTDSJCFE CFMPX JT FBTJMZ BTTFNCMFE XPSLT BT B DMBTT EFNPOTUSBUJPO TQBXOT TNBMM HSPVQ QSPKFDUT GJUTBWBSJFUZPGDMBTTUJNFGSBNFTJODMVEJOHMBCTBOEQSPWJEFTGPS TVCTUBOUJBMEJTDVTTJPO .FUIPET 5IJTFYFSDJTFSFRVJSFTMJUUMFNPSFUIBOBCJUPGDSFBUJWFOFTT BOEBUSJQUPUIFHSPDFSZTUPSF'PSBDMBTTSPPNEFNPOTUSB UJPOXFTVHHFTUUIBUZPVDPOEVDUUIJTFYFSDJTFJOBTFUUJOHUIBU BMMPXTBDDFTTUPBMBSHFXSJUJOHTVSGBDFXIJUFCPBSECMBDLCPBSE PSQPTUFST 'PSBNPSFUJNFDPOTVNJOHBOEJOUFOTJWFJOWFTUJHB UJPOTVDIBTBMBCQFSJPE USJBMWFSTJPOTPGQIZMPHFOFUJDBOBMZTJT QSPHSBNT TVDI BT .BD$MBEF IUUQNBDDMBEFPSHNBDDMBEF IUNM DBOBMTPQSPWFVTFGVMGPSHFOFSBUJOHDMBEPHSBNTCBTFEPO UIFDIBSBDUFSTDIPTFOCZZPVSDMBTT'JHVSF 5IFCBTJDJEFBPG UIJT FYFSDJTF DPVME CF FYFDVUFE XJUI BOZ OVNCFS PG FYBNQMFT XIJDIFOBCMFTJUUPCFBEBQUFEUPTVJUPOFµTPXOJOUFSFTU 5P GPMMPX PVS FYBNQMF QVSDIBTF UIF GPMMPXJOH DBOEJFT 232 #BCZ3VUI#3 )&34)&:µ4DIPDPMBUFCBS)$ )&34)&:µ4XJUI"MNPOET)" )&34)&:µ48IJUF$IPDPMBUF)8$XIJDITPNFUJNFT DPNFTXJUI$PPLJFTOµ$SFBN THE AMERICAN BIOLOGY TEACHER, VOLUME 69, NO. 4, APRIL 2007 ..µ41MBJO..1- ..µ41FBOVU..1& 3&&4&µ41*&$&431 4/*$,&344/ 4/*$,&34"MNPOE#BS4/" 4/*$,&34$SVODIFS4/$ 5IF BEWBODFE DSJUJDBM UIJOLJOH FYFSDJTFT BMTP SFRVJSF 3&&4&µ4 QFBOVU CVUUFS DVQT 31#$ BOE 4,*55-&4 4, 5IFBCCSFWJBUJPOTGPSUIFTFDBOEJFTBQQFBSPOUIF EBUBNBUSJYBOETPNFDMBEPHSBNT/PUFGPSHSPVQFYFS DJTFTUIBUNVMUJQMFTFUTPGUIFDBOEJFTXJMMCFOFFEFE 'JSTU JOUSPEVDF TUVEFOUT UP UIF HPBM PG QIZMPHFOFUJD BOBMZTJT JF EFUFSNJOJOH UIF NPTU QBSTJNPOJPVT USFF 1SFGBDF UIF FYFSDJTF XJUI BO JOUSPEVDUPSZ EJTDVTTJPO PG IPX TDJFOUJTUT USZ UP EFUFSNJOF UIF QIZMPHFOFUJD SFMB UJPOTIJQT CFUXFFO PSHBOJTNT BOE UIF JNQMJDBUJPOT UIJT IBTGPSUIFTUVEZPGFWPMVUJPO4FWFSBMRVBMJUZ8FCTJUFT 6OEB /$#* 5SFF PG -JGF 1SPKFDU DBOQSPWJEFUIFJOTUSVDUPSXJUICBDLHSPVOEJOGPSNBUJPO %JTUJOHVJTI CFUXFFO QIZMPHFOZ DMBEJTUJDT BOE DMBT TJGJDBUJPOBTEFTDSJCFEBCPWF3FNJOEUIFTUVEFOUTUIBU BMM DMBEPHSBNT BSF UZQFT PG QIZMPHFOFUJD USFFT CVU OPU BMMQIZMPHFOFUJDUSFFTOFFECFDMBEPHSBNT&NQIBTJ[FUP UIFTUVEFOUTUIBUUIFZBSFOPUDMBTTJGZJOHUBYBCVUJOTUFBE HFOFSBUJOHIZQPUIFTFTJFDMBEPHSBNT BCPVUQIZMPHF OFUJDSFMBUJPOTIJQT /FYU HJWF UIF TUVEFOUT UIF UXP QBDLBHFT PG . .µ4 1MBJOBOE1FBOVU BOEUIFQBDLBHFPG3&&4&µ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µ4 1FBOVU #VUUFS MPPL TJNJMBS UP 3&&4&µ4 1*&$&4 JO WBMJE DIBSBDUFST DBOEZ DPBU JOH QFBOVU CVUUFS CVU POMZ UIF ..µ4SFUBJODIPDPMBUF5IFO BGUFS UIFZ IBWF TVDDFTTGVMMZ DPO TUSVDUFE POF QPTTJCMF DMBEPHSBN SFRVFTUUIBUUIFTUVEFOUTJEFOUJGZ UIF TZOBQPNPSQIJFT PS TIBSFE DIBSBDUFSJTUJDT JF JOEJWJEVBM QJFDFTDBOEZDPBUJOH BOETQFDV MBUF BCPVU UIF DPNNPO BODFTUPS BUFBDICSBODIPGUIFUSFF %FQFOEJOH PO IPX RVJDLMZ UIF TUVEFOUT HSBTQ UIF JEFBT ZPVNBZBMTPTVHHFTUUIBUUIFZ CVJMEBTFQBSBUFDMBEPHSBNGPS UIPTF DBOEJFT UIBU DPNF JO CBS GPSN 'JHVSF # 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Two most parsimonious cladograms, each with 13 steps. A) Common ancestor assumed to be the chocolate bar B) Common ancestor assumed to be candy coated chocolate pieces. Perpendicular lines on branches represent evolutionary steps and text defines addition or loss of a character. Parsimonious phylogenies *O .BD$MBEF UIJT QSPDFTT PG BSSJWJOH BU UIF NPTU QBSTJNPOJPVT USFF JT EPOF CZ NPWJOH UIF CSBODIFT PGBEFGBVMUDMBEPHSBNVTJOHUIFBSSPXUPPM'JHVSF VOUJM ZPV IBWF NJOJNJ[FE UIF USFF MFOHUI CBTJOH UIJT OVNCFS GSPN UIF DIBSU ZPV QSPEVDFE FBSMJFS +VTU BT ZPV XPVME EP PO QBQFS ZPV TIPVME NPWF UIF CSBODIFT UP QPTJUJPOT UIBU NBLF TFOTF JO MJHIU PG BOZ IJTUPSJDBM SFDPSET UIBU ZPV NBZ IBWF 'JHVSF *OUIFGVMMWFSTJPOJUJTQPTTJCMFUPVTF1"61 1IZMPHFOFUJD"OBMZTJT6TJOH1BSTJNPOZIUUQQBVQ DTJUGTVFEVBCPVUIUNM UPIFMQJEFOUJGZUIFNPTUQBS TJNPOJPVTUSFF 3FTVMUT%JTDVTTJPO 'PSPVSFYBNQMFXFEJTDVTTUXPQBSTJNPOJPVTDMBEPHSBNT FBDIXJUITUFQT'JHVSF"# UIBUXFSPPUFEJOEJGGFSFOUPVU HSPVQT3FHBSEMFTTPGXIFSFXFSPPUUIFUSFFFJUIFSXJUIBDMBTTJD )&34)&:µ4CBSPSXJUIDBOEZJOQJFDFTUIFNPTUQBSTJNPOJPVT CSBODIJOHJFUPQPMPHZ SFNBJOTUIFTBNF5IFUXPUSFFTJMMVT USBUFUIBUUIFEJGGFSFODFDPNFTXJUIUIFJOGFSFODFUPUIFEJSFDUJPO PGDIBOHFJFMPTTPSHBJOPGBHJWFODIBSBDUFS*O'JHVSF"DBOEZ DPBUJOHJTBEEFEBOEUIFCBSGPSNMPTUUPBSSJWFBUUIFDBOEJFTXJUI QJFDFT*GZPVTUBSUXJUIQJFDFTJOUIFCPUUPNDMBEPHSBN'JHVSF # UIFO UIF DBOEZ DPBUJOH JT MPTU BOE UIF CBS GPSN HBJOFE UP DSFBUF UIF )&34)&:µ4 DMBEF 4UVEFOUT DBO GVSUIFS FYQMPSF UIF EJSFDUJPOBMJUZ PG DIBOHF CZ FYBNJOJOH BOZ BTTVNQUJPOT UIBU UIFZ NBEF JO DPOTUSVDUJOH UIFJS DMBEPHSBNT *O PVS FYBNQMF XF BTTVNFE UIBU QFBOVU CVUUFS DBNF GSPN QFBOVUT CBTFE PO PVS SFTFBSDI 'JHVSF BOE FBSMZ TDIPPM MFTTPOT BCPVU (FPSHF 8BTIJOHUPO$BSWFS)PXFWFSJOCVJMEJOHQIZMPHFOFUJDUSFFTJUJT KVTUBTQBSTJNPOJPVTGPSUIFDIBOHFUPHPJOUIFPUIFSEJSFDUJPO 234 THE AMERICAN BIOLOGY TEACHER, VOLUME 69, NO. 4, APRIL 2007 XIFSFQFBOVUTDPBMFTDFPVUPGBTPGUBNBMHBNPG²QFBOVUUZ³NBUF SJBM,OPXJOHUIFBODFTUSBMDPOEJUJPOCRTKQTKBMMPXTVTUPNBLF CFUUFSFEVDBUFEHVFTTFTBTUPUIFNPTUQMBVTJCMFQIZMPHFOZ 'JHVSF"VTFT)&34)&:µ4DIPDPMBUFCBSBTUIFPVUHSPVQ #BTFEPOUIFIJTUPSZPGDIPDPMBUF'JHVSF XFLOPXUIBU)$ BQQFBSFECFGPSFBOZPGUIFPUIFSDBOEJFTBOEJTUIFMPHJDBMDIPJDF GPSUIFPVUHSPVQ5IFFWPMVUJPOBSZDPODMVTJPOGSPNUIJTJTUIBU UIFPVUHSPVQCSBODIFEGSPNUIFQBSFOUHSPVQCFGPSFUIFPUIFS HSPVQTPSDMBEFT CSBODIFEGSPNFBDIPUIFS)PXFWFSXJUIPVU UIJT LOPXMFEHF TUVEFOUT DPVME NBLF DPOWJODJOH BSHVNFOUT GPS UIF SFWFSTF TDFOBSJP UIBU CFHJOT XJUI DBOEZ DPBUFE QJFDFT *OTUSVDUPSTDPVMEBTLTUVEFOUTUPEFGFOEUIFJSDIPJDFTBOEUIFO JOUSPEVDF UIF ²IJTUPSJDBM FWJEFODF³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²MJNJUT³ PG UIJT FYFSDJTF POF DPVME JODMVEF B EJTDVTTJPO PGBSUJGJDJBMTFMFDUJPOJFUIFTFMFDUJWFCSFFEJOHPGEPNFTUJDBUFE QMBOUT BOE BOJNBMT UP FODPVSBHF UIF PDDVSSFODF PG EFTJSBCMF USBJUT0GDPVSTFDBOEZCBSTEPOPUOBUVSBMMZSFQSPEVDFCVUUIF QPJOUJTUIBUIVNBOTCZUIFJSCVZJOHIBCJUTTFMFDUUIFTFUSBJUT BSUJGJDJBMMZBTXFIBWFEPOFXJUIPVSDSPQTMJWFTUPDLBOEQFUT*O PVSDMBEPHSBNTXFBMTPTFFFYBNQMFTPGIPNPQMBTZXIFSFTJNJ MBSJUZPDDVSTJOEFQFOEFOUMZ PG FWPMVUJPOBSZ DIBOHF -PTT PG DIPDPMBUF UP IBWF FJUIFS 3&&4&µ4 1*&$&4 PS )&34)&:µ4 8IJUF $IPDPMBUF XPVME CF POF FYBNQMF *O BEEJUJPO UIF BSSBOHFNFOUPGUIFJMMVTUSB UJPOT JOUP DMBEFT FNQIB TJ[FT UIF TJTUFS HSPVQT UIBU PDDVS JODMVEJOH . .µ4 4/*$,&34 WBSJFU JFTBOE)&34)&:µ4CBST 5IFSFGPSF UIJT FYFSDJTF QSPWJEFT PQQPSUVOJUJFT UP EJTDVTT UIF JOUSJDBDJFT UIBU %BSXJO PCTFSWFE XIFO IF GJSTU EFTDSJCFE EFTDFOU XJUINPEJGJDBUJPO Figure 7. Change in data matrices and cladograms with additions of new taxa. A) REESE’S peanut butter cups B) SKITTLES®. A non-shaded rectangular box shows score of new taxon in the data matrix and a circle shows the location in the cladogram. 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