darwin and evolution: a set of activities based on the...

SSR June2010,91(337) 75

Darwin and evolution: a set of activities based on the evolution of mammals

Janet M. Haresnape

ABSTRACT These activities, prepared for key stage 5 students (ages 16–18) and also suitable for key stage 4 (ages 14–16), show that physical appearance is not necessarily the best way to classify mammals. DNA structure is examined to show how similarities and differences between DNA sequences of mammals can be used to establish evolutionary relationships. Some real DNA sequences are compared to illustrate that the whale is surprisingly closely related to mammals such as the hippo, camel and cow, a relationship confirmed by fossil evidence. Finally, an analysis of haemoglobin sequences of primates shows the close relationship between humans and other great apes such as the chimpanzee.

Darwinmadeextensiveobservationsonagreatmanycreatures,includingmammals,andnoticedthatspeciesfellintonaturalgroups.Forexample,lions,tigersandleopardshavemanysimilaritiesandresemblecats.Onthebasisofhisobservations,hewasabletoplacemammalsindistinctgroups.Hisworkhasbeencontinued,andwenowrecognisethatallmammalsevolvedfromacommonancestorandhavebranchedintomanydifferenttaxonomicorders(Figure1).

Theactivitiesdescribedhereprovidestudentswithanopportunitytocomparespeciesonananatomicalandbiochemicallevel.InformationavailabletoDarwin,suchasfossilevidenceandobservationofthephysicalappearanceofanimals,isconsideredalongsidethemolecularevidencethatisavailablenow.StudentsexaminegenuinemolecularevidenceintheformofDNAandaminoacidsequences,andusethistodeduceevolutionaryrelationshipsbetweendifferentmammals.Theimportanceoflookingatevidencefromdifferentsources,suchasmorphological,fossilandmolecular,isemphasised.Theseproblem-solvingexercisesallowsmallgroupsofstudentstothinkthroughtheevolutionaryrelationshipsbetweenmammaliangroups,andtorealisethatphysicalappearanceisnotalwaysthebestwaytoclassifymammals.Theactivitieshavebeensuccessfullytrialledwithagroupofyear9students(ages13–14)aswellaswithkeystage5(ages16–18)studentsforwhomtheywere

prepared.Theytakeapproximatelytwohourstocomplete,butthiscouldbespreadovertwoormorelessons.Whereappropriate‘blank’versionsoftablesandfiguresareprovidedwhichcanbecopiedforstudentstocomplete.

Activity 1: Odd one out

Thefirstactivityinvolvesanexaminationoffourwell-knownmammals:therhinoceros,hippopotamus,elephantandwhale.

Webeginbyaskingwhichofthefouristhe‘oddoneout’.Modelsorpicturesofthesefourmammalsshouldbepresentedifpossible,andstudentsaskedtoconsiderwhichoneappearsatfirstsighttobemostdifferentfromtheotherthree.

Studentsarelikelytosuggestthatthewhaleistheoddoneoutonthebasisofitshabitat,asitlivesinthesea,andalsoonthebasisofitsappearance,asithasnolegs.Othersmaysuggesttherhino,asitisthemostendangered,ortheelephant,asithasatrunk.Agoodargumentcanbemadeforanyanswer.However,inevolutionaryterms,itistheelephantthatismostdifferentfromtheothers,asithasadifferentevolutionaryoriginfromtheotherthree.(Itisinthepurplesection,inFigure1,whereastherhino,hippoandwhaleareallinthebluesection.)Thephysicalappearanceofanimalsdoesnotnecessarilyindicatewhicharemostcloselyrelated.

Wenowturntolookatsomefossilevidencethattellsusabouttheevolutionaryhistory

76 SSRJune2010,91(337)

Darwin and evolution: a set of activities based on the evolution of mammals Haresnape

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SSR June2010,91(337) 77

ofmodernwhales.Figure2summarisestheevolutionaryoriginofwhalesanddolphins,andshowsthattheyevolvedfromlandcreatureswithfourlegs,graduallylosingtheirlegsandbecomingmorestreamlined.Notethatfossilsofancestral

whaleshaveananklebone,eventhoughtheanimalhadnolegs,showingthatitwasdescendedfromanimalswhichdidhavelegs.ThisisclearlyvisibleinthefullyaquaticDorudon(Figure2)andcanstillbeseenintheskeletonsofmodernwhales.

Figure 2 Somerepresentativemammalsintheevolutionaryhistoryofmodernwhales.Imaginativereconstructionsofthefossilspeciesareshown,suggestinghowtheymighthavelookedinlife.Notethesearenotdrawntoscale,andthemeasurementsrefertobodylength.(Figure7.5fromThomas,2003,reproducedwithgratefulpermission.)

Haresnape Darwin and evolution: a set of activities based on the evolution of mammals

78 SSRJune2010,91(337)

Thisfossilevidence,togetherwiththeevolutionarytree(Figure1),showsthatthewhaleismorecloselyrelatedtothehippothanthehippoistotheelephant,oreventherhino,despiteitsverydifferentoutwardappearance.

Activity 2: Spot the difference

Wenowturntolookathowmolecularevidencecanbeusedtoinvestigateevolutionaryrelationships.Thisactivityinvolvesexaminingtwomodelsofdouble-strandedDNA,withonedifferenceinbasesequencebetweenthem.Thisillustrateswhatismeantbyamutation,ordifferenceinbasesequence.Ifmodelsarenotavailable,adiagramsuchasthatshowninFigure3canbeused.Thisactivityprovidesanopportunitytodiscusscomplementarybasepairingindouble-strandedDNA,andthefactthatcloselyrelatedspecieshavesimilarDNA.ThenumberofmutationsordifferencesbetweentheDNAofdifferentspeciescangiveanindicationofhowlongagotheydivergedfromacommonancestor.

Activity 3: DNA sequence evidence

Table1showstheDNAsequenceofasmallstretchofthegeneforaparticularmilkproteincalledcaseininanumberofdifferentmammals.Thesequencediffersfromonemammaltoanother,andthoseshownarefromtwospeciesofwhale,severalfamiliarungulatespecies(membersoftheorderArtiodactyla,includinghippo,pig,cow,sheep,goat,deerandcamel,pluszebraandtapirfromtheorderPerissodactyla),andtheleopard(Carnivora).AlltheseareinthebluesectionoftheevolutionarytreeshowninFigure1.Alsoincludedaresomemoredistantlyrelatedmammals,fromordersinthegreensectionofthetree:human,rabbitandmouse.Ideally,thesesequenceswouldbepresentedasstringsofcolouredbeads,withonecolourrepresentingeachofthefourbases,A,T,CandG.Alternatively,Table1couldbeprintedincolourandcutintostripsforcomparisonofsequencesfromdifferentspecies.

ThetaskistocounthowmanydifferencesthereareinthisstretchofDNAbetweenthewhaleandeachoftheothermammals(studentscanbegivenacopyofTable2atocomplete).Ifthesequencesarerepresentedasstringsofbeads,orasstripsofpaper,thecalculationcanbedonefairlyeasilybyliningthemupagainsteachotherandcountingthenumberofcolourdifferences.ThedifferencesareshowninthelastcolumnofTable2b,andconfirmthatwhaleDNAcloselyresemblesthatofthemembersoftheorderArtiodactylaandthatthewhaleisthereforecloselyrelatedtothem.Herewhalesappeartobemostcloselyrelatedtothehippo,coworcamel,asthereareonlythreedifferencesbetweentheirbasesequencesinthisstretchofDNA.

NotethatthedifferencebetweenthewhaleDNAandthatofthehuman,rabbitormouseismuchlargerthanbetweenthewhaleandothermammalsinthetable.Thesemammalsaremoredistantlyrelated,beinginthegreensectionasopposedtothebluesectionoftheevolutionarytreeshowninFigure1.

Moredetailedstudiesofthiskind,lookingatmuchlongersequencesfrommanydifferentgenes,haveconfirmedthatwhalesanddolphinsdohavetheirevolutionaryoriginwithintheArtiodactyla,andthisisinagreementwiththeevidencefromfossils.Theirclosestlivingrelativeisactuallybelievedtobethehippo.

Astutestudentsmaynotewithsurprisethatthenumberofdifferencesbetweenthewhaleandthe


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Figure 3 Twoshortsequencesofdouble-strandedDNAwithjustonebasedifference.BasesareA(adenine),C(cytosine),G(guanine)andT(thymine).Inthefifthbasepairfromthetop,AhasbeenreplacedbyTandThasbeenreplacedbyA.

SSR June2010,91(337) 79

Table 1 DNAsequencesfrompartofthecaseingeneofselectedmammals.Thisisa50basesequencetakenfromexon7ofthegeneforbeta-casein.BasesareA(adenine),C(cytosine),G(guanine)andT(thymine).(FromGatesyet al.,1996.)

Toot

hed

wha

le

Bal

een

wha

le

Hip

popo

tam

us

Cow

Cam

el

Gira

ffe

She

ep

Goa

t

Pec

cary

Pig

Dee

r

Tapi

r

Zebr

a

Leop

ard

Hum

an

Rab

bit

Mou

se

1 A A A T A T T T A A T A A A G G G 2 G G G G G G G G G G G G G G A G C 3 A A A A T A A A A A A T T T T T T 4 G G G G G G G G G G G G G G A C C 5 C C C C C C C C C C C C C C C C C 6 C C C C C C C C C C C C C C C C G 7 C C C C C C C C T T C C C C C C C 8 T T T T T T T T T T T T T C T T C 9 T T T T T T T T T T T T T T T T T10 T T T T T T T T T T C T T T T T C11 A A A A A A A A G G G T T T T G A12 T T C C C C C C T T C C C T T T T13 T T T T T T T T T G T T T G T T A14 G G G G G G G G G G G G G G G G A15 A A A A A A A A A A A A A A A A A16 A A A A A A A A A A A A A C C C C17 A A A A A A A A G G G C C A C T T18 G G G G G G G G G G G G G G C C C19 C C G C C C C C C C C C C C T T T20 C C C C C C C C C C C C C C C C C21 A A A A A A A A A A A A A A A A A22 G G G G G G G G G G G A A A A A A23 A A A A A A A A A A A A A A A A A24 G G G G G G G G G G G T T T T T T25 C C C C C C C C C C C C C C C T C26 C C C C C C C C C C C T T G C C C27 T T T T T T T T T T T T T T C T C28 G G G G G G G G A A G G G G A G C29 A A A A A A A A A A A A A A A A A30 C C C C C C C C C C C A A A A A G31 T T T T C T T T T T T T T T A T C32 C C C C C C C C C C C C C C C C C33 T T T T T T T T T T T C C T T T T34 C C T C G C C C C C C C C C C C T35 A A A A A A A A A A A A A A A A G36 C C C C C C C C C C C C C C C G C37 T T T T T T T T T T T T T T T G C38 G G G G G G G G G G G A A G G G A39 A A A A A A A A A A A A A A A A G40 T T T T T T T T T T T T T T T G T41 C C C G C G G G T T G G G C C A C42 T T T T T C T T T T T G G T T T T43 T T T T C T T T C T T T T T T G T44 G G G G G G G G G G G G G G G A G45 A A A A A A A A A A A A A A A A C46 A A A A A A A A A A A A A A A A T47 A A A A A A A A A G A A A A A A A48 A A A A A A A A G T A A A A A A A49 T T T T T T G G T C C T T T T T T50 C C C C C C C C C C C C C C C C C


80 SSRJune2010,91(337)

leopardislessthanbetweenthewhaleandzebraortapir.ThisisbecausethelengthofDNAunderconsiderationisonly50baseslong,andanalysisoflongersequenceshasestablishedthattherelationshipsshowninFigure1arecorrect.

Iftimepermits,amoredetailedanalysisofthesequencesshowninTable1couldbeundertaken;forexample,comparingmoreoftheDNAsequencesinapairwisemanner.ApossiblecomparisoncouldinvolvethemammalsshowninTable3.Ofthese,thesheepandgoatformthemostcloselyrelatedpair,withnoDNAsequencedifferencesinthisparticular50basestretchofDNA.ThezebraismoredistantlyrelatedthanthemammalswithintheorderArtiodactyla,aswouldbeexpectedasitisamemberoftheorderPerissodactyla(Figure1).TheDNAevidenceinTable3bconfirmsthatthewhaleismorecloselyrelatedtotheArtiodactylaspeciesthananyofthosewithintheorderArtiodactylaaretothezebra.Iftimeisshort,itmightbemoreworthwhiletoomitthisanalysisandmoveontoActivity4,whichisasimilarproblem-solvingexercise.

Activity 4: Haemoglobin sequences of primates

Herewetakesevenspeciesofprimateandconsiderourrelationshiptootherprimategroupsbycomparingthesequenceofthefirst130aminoacidsofthebeta-haemoglobinchainfoundineachofthem.Inadditiontohumans,weconsiderthegreatapes(chimpanzeeandgorilla),alesserape(gibbon),anOldWorldmonkey(rhesusmonkey),aNewWorldmonkey(squirrelmonkey)andalemur.Studentsmightfirstbeaskedtopredictwhichoftheseprimatespeciesaremostcloselyandmostdistantlyrelatedonthebasisoftheirappearance.Picturesormodelsofthespeciescouldbeprovided.Itislikelythattheywillsuggestthatthetwospeciesofmonkey(rhesusmonkeyandsquirrelmonkey)aremorecloselyrelatedthanthedifferentspeciesofgreatape(gorilla,chimpanzee,human).

Afterconsiderationofthisquestion,studentscouldbeshownthehaemoglobinsequencesofthesevenprimates,showninTable4.Thoseaminoacidswhicharethesameforallprimatespeciesareshowningrey,soonlythosethatdifferareidentifiedhere.Eachaminoacidinthepositionswherethereisvariationisrepresentedbyadifferentcolour,identifiedinthekey.Again,these


Table 2 DNAsequencedifferencesbetweenthewhaleandeachoftheotherspeciesforthestretchofthegeneshowninTable1.(a)Forstudentstocomplete,(b)completed.

(a)

Mammal species

Order Number of differences in this stretch of DNA be-tween this species and the whale

Toothedwhale CetaceaBaleenwhale CetaceaHippopotamus ArtiodactylaCow ArtiodactylaCamel ArtiodactylaGiraffe ArtiodactylaSheep ArtiodactylaGoat ArtiodactylaPeccary ArtiodactylaPig ArtiodactylaDeer ArtiodactylaTapir PerissodactylaZebra PerissodactylaLeopard CarnivoraHuman PrimatesRabbit LagomorphaMouse Rodentia

(b)

Mammal species

Order Number of differences in this stretch of DNA be-tween this species and the whale

Toothedwhale Cetacea 0Baleenwhale Cetacea 0Hippopotamus Artiodactyla 3Cow Artiodactyla 3Camel Artiodactyla 3Giraffe Artiodactyla 4Sheep Artiodactyla 4Goat Artiodactyla 4Peccary Artiodactyla 5Pig Artiodactyla 7Deer Artiodactyla 7Tapir Perissodactyla 11Zebra Perissodactyla 11Leopard Carnivora 9Human Primates 15Rabbit Lagomorpha 18Mouse Rodentia 25

SSR June2010,91(337) 81

sequencesarebestshownasstringsofcolouredbeads,withgreybeadsfortheinvariantsites,andadifferentcolourforeachaminoacid,inordertomakethecomparisonseasier.However,asinActivity3,itwouldbepossibletoprintacolourversionofTable4andcutitintostripsforpairwisecomparisonofspecies.

AswithDNAsequences,analysisofthesimilaritiesanddifferencesbetweentheaminoacidsequencesinthesameproteinfromdifferentmammalscangiveanindicationofhowcloselyrelatedtheyare.MoststrikinginTable4arethesimilaritiesbetweentheaminoacidsequencesofbeta-haemoglobininallprimates,shownbyallthepositionscolouredingrey.Ninety-fiveofthe130aminoacidsinthispartofthechain(73%)arethesameforallprimates.

Detailedanalysisofthedifferencesinaminoacidsequencecanbemadebycountingthe

differencesbetweenpairsofprimatespeciesandrecordingtheminacopyofTable5a.Table5bisacompletedversionofthetable.Ifthereisinsufficienttimeforthis,studentscouldsimplylookatthenumberofdifferencesbetweenthehumansequenceandeachoftheotherprimates.Eitherway,itisclearfromthisevidencethathumansareverycloselyrelatedtothegreatapes,suchasthegorillaandchimpanzee.Theaminoacidsequencesofbeta-haemoglobininhumansandchimpanzeesareidentical,andthereisonlyonedifferencebetweenthesequencesinhumansandgorillas.

FromacompletedversionofTable5,theevolutionaryrelationshipsbetweenthesesevenspeciesofprimatecanbededuced,andareshowninFigure4.ItisworthpointingoutthatthenumberofdifferencesbetweenthesequenceofanOldWorldmonkey(rhesusmonkey)andthatofaNewWorldmonkey(squirrelmonkey)

Table 3 PairwisecomparisonofDNAsequencedifferencesbetweenthewhale,fivespeciesfromtheorderArtiodactyla(hippo,cow,camel,sheepandgoat)andthezebrafromtheorderPerissodactyla.(a)Forstudentstocomplete,(b)completed.

(a)NumberofDNAsequencedifferencesbetweenpairsofmammals

Whale Hippo Cow Camel Sheep Goat Zebra

Whale

Hippo

Cow

Camel

Sheep

Goat

Pig

(b)NumberofDNAsequencedifferencesbetweenpairsofmammals(completed)

Whale Hippo Cow Camel Sheep Goat Zebra

Whale 3 3 3 4 4 12

Hippo 4 4 5 5 13

Cow 4 1 1 11

Camel 5 5 11

Sheep 0 11

Goat 11

Pig


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Table 4 Aminoacidsequenceofbeta-haemoglobininsevenspeciesofprimate.Thesecouldbepresentedinanyorder,butareheregiveninorderoftheclosenessoftheirrelationshiptohumans.Greypositionsrepresentthoseaminoacidswhicharethesameforalltheseprimates.(AdaptedfromBanister-Marx,J.TheNationalHealthMuseumactivitiesexchange.)

Positioninchain

Primatespecies

Hum

an

Chi

mp

Gor

illa

Gib

bon

Rhe

sus

mon

key

Squ

irrel

m

onke

y

Lem

ur1 VAL VAL VAL VAL VAL VAL THR2 HIS HIS HIS HIS HIS HIS LEU34 THR THR THR THR THR THR SER5 PRO PRO PRO PRO PRO GLY ALA6 GLU GLU GLU GLU GLU ASP GLU78 LYS LYS LYS LYS LYS LYS ASP9 SER SER SER SER ASN ALA ALA

10 ALA ALA ALA ALA ALA ALA HIS1112 THR THR THR THR THR ALA THR13 ALA ALA ALA ALA THR ALA SER1415161718192021 ASP ASP ASP ASP ASP GLU GLU22 GLU GLU GLU GLU LEU ASP LYS2324252627282930313233 VAL VAL VAL VAL LEU VAL VAL3435363738

Positioninchain

Primatespecies

Hum

an

Chi

mp

Gor

illa

Gib

bon

Rhe

sus

mon

key

Squ

irrel

m

onke

y

Lem

ur

394041424344454647484950 THR THR THR THR SER THR SER5152 ASP ASP ASP ASP ASP ASP SER53545556 GLY GLY GLY GLY GLY ASN SER57585960616263646566676869 GLY GLY GLY GLY GLY GLY SER70717273 ASP ASP ASP ASP ASP ASP GLU747576 ALA ALA ALA ALA ASN THR HIS77787980 ASN ASN ASN ASP ASN ASN ASN81828384


SSR June2010,91(337) 83

Positioninchain

Primatespecies

Hum

an

Chi

mp

Gor

illa

Gib

bon

Rhe

sus

mon

key

Squ

irrel

m

onke

y

Lem

ur

858687 THR THR THR LYS GLN GLN GLN888990919293949596979899

100101102103104 ARG ARG LEU ARG LYS ARG LYS105106107108109110111 VAL VAL VAL VAL VAL VAL SER112 CYS CYS CYS CYS CYS CYS ALA113 VAL VAL VAL VAL VAL VAL GLU114 LEU LEU LEU LEU LEU LEU SER115 ALA ALA ALA ALA ALA ALA GLU116 HIS HIS HIS HIS HIS HIS LEU117118119120 LYS LYS LYS LYS LYS LYS HIS121 GLU GLU GLU GLU GLU GLU ASP122 PHE PHE PHE PHE PHE PHE LYS123 THR THR THR THR THR THR SER124125 PRO PRO PRO GLN GLN GLN ALA126 VAL VAL VAL VAL VAL LEU VAL127128129130 TYR TYR TYR TYR TYR TYR PHE

KeytoaminoacidsinTable4

Abbreviation Aminoacid

ALA alanine

ASN asparagine

ARG arginine

ASP asparticacid

CYS cysteine

GLN glutamine

GLY glycine

GLU glutamicacid

HIS histidine

LEU leucine

LYS lysine

PHE phenylalanine

PRO proline

SER serine

TYR tyrosine

THRTTHR threonine

VAL valine


84 SSRJune2010,91(337)

ismorethanbetweenhumansandapes.Thisisbecausethesetwogroupsofmonkeysdivergedlongeragothanhumansdivergedfromotherapes.Soalthoughtheyseemmoresimilarinoutwardappearancethanhumansandothergreatapes,theyaremuchmoredistantlyrelated.Byexaminingevidencefromfossils,andmolecularandanatomicaldata,ithasbeendeducedthathumanandchimpanzeelineagesdivergedabout6millionyearsago,whereasOldWorldandNewWorldmonkeysdivergedabout13millionyearsago.

SimilaraminoacidsequencesofaproteinindicatethatsimilarDNAsequencescodefor

them.Overall,99%oftheDNAofhumansandchimpanzeesisidentical.The1%oftheDNAwhichisdifferentiscrucialindeterminingwhatmakesushuman,andaccountsforallthedifferencesbetweenhumansandchimpanzees.Theremaining99%isresponsibleforalltheattributesofchimpanzeeswhicharesimilartothoseofhumans.Wearemuchmoresimilartochimpsthanwemightthink.

Toseehowyoumighthavelookedmillionsofyearsago,gotowww.openuniversity.co.uk/darwinandmorphyourownfaceusingtheOpenUniversity’sfunfacemorphtool,‘Devolveme’.

Table 5 Numberofaminoaciddifferencesbetweendifferentpairsofprimates.(a)Forstudentstocomplete,(b)completed.

(a)Numberofaminoaciddifferencesbetweenbeta-haemoglobinchains

Human Chimp Gorilla Gibbon Rhesusmonkey

Squirrelmonkey Lemur

Human

Chimp

Gorilla

Gibbon

Rhesusmonkey

Squirrelmonkey

Lemur

(b)Numberofaminoaciddifferencesbetweenbeta-haemoglobinchains(completed)

Human Chimp Gorilla Gibbon Rhesusmonkey

Squirrelmonkey Lemur

Human 0 1 3 10 10 30

Chimp 1 3 10 10 30

Gorilla 4 10 12 30

Gibbon 9 11 31

Rhesusmonkey 13 29

Squirrelmonkey 30

Lemur


SSR June2010,91(337) 85

SomeoftheseactivitiesareavailableonlineintheScienceandNaturesectionoftheOpenUniversity’sOpenLearnwebsite(Haresnape,2010).

AcknowledgementsThankstoMartinScottandLindaHurst(Open

UniversityintheSouth)forhelpwithpreparingFigure1,andtotherefereesfortheirhelpfulcomments.ThisworkwasdevelopedattheOpenUniversityandhasbeenadaptedforSchool Science Reviewwithpermission.

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Figure 4 Evolutionaryrelationshipsbetweenprimategroups,asdeducedfromaminoacidsequenceevidence.(a)Forstudentstocomplete,(b)completed.


86 SSRJune2010,91(337)

References

Banister-Marx,J.TheNationalHealthMuseumactivitiesexchange.Molecular biology and primate phylogenetics.www.accessexcellence.org/AE/AEPC/WWC/1995/simulation_molecular.php(accessedJuly2009).

Gatesy,J.,Hayashi,C.,Cronin,M.A.andArctander,P.(1996)Evidencefrommilkcaseingenesthatcetaceansarecloserelativesofhippopotamidactiodactyls.Molecular Biology and Evolution,13,954–963.

Haresnape,J.M.(2010)TheOpenUniversity’sOpenLearn

website.Schoolactivities:Evolutionary tree of mammals.Availableat:openlearn.open.ac.uk

MacDonald,D.ed.(2001)What is a mammal? The new encyclopedia of mammals.Oxford:OxfordUniversityPress.

Thomas,J.ed.(2003)Studying mammals.OpenUniversityScienceShortCourse,S182.MiltonKeynes:OpenUniversity.

Janet Haresnape isfromtheLifeSciencesDepartmentoftheOpenUniversity,andhasbeenamajorcontributortotheproductionanddeliveryoftheOpenUniversityEvolutioncourse(S366).Email:[email protected]


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