Index to this page

-atural (election The .easure o# /!itness/ o (ur$i$al o (e)ual (election o !ecundit& (election 0re %umans E)empt #rom -atural (election1 Continuous 2ariation %eritabilit& The E##ects o# (election on 3opulations o (tabili4ing (election o Directional (election o Industrial .elanism
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Disrupti$e (election

Evolution and Adaptation

In 1859 the English naturalist Charles Darwin published The Origin of Species. The book contained two major arguments !irst" Darwin presented a wealth o# e$idence o# evolution. %e said that all li$ing things on earth toda& are the descendants ' with modi#ications ' o# earlier species. (econd" he proposed a mechanism ' natural selection ' to e)plain how e$olution takes place. E$olution in$ol$es two interrelated phenomena adaptation *$er the course o# time" species modi#& their phenot&pes in wa&s that permit them to succeed in their en$ironment. This page is de$oted to looking at how e$olution leads to adaptation.

speciation *$er the course o# time" the number o# species multiplies+ that is" a single species can gi$e rise to two or more descendant species. In #act" Darwin maintained that all species are related+ that is" an& two species on earth toda& ha$e shared a common ancestor at some point in their histor&. The process o# speciation is e)amined in a separate page. ,ink to it.

Natural Selection

,i$ing things produce more o##spring than the #inite resources a$ailable to them can support. Thus li$ing things #ace a constant struggle for existence. The indi$iduals in a population $ar& in their phenotypes. (ome o# this $ariation is inheritable+ that is" it is a re#lection o# $ariations in genotype. Those $ariants best adapted to the conditions o# their li#e are most likel& to sur$i$e and reproduce themsel$es 5/sur$i$al o# the #ittest/6. To the e)tent that their adaptations are inheritable" the& will be passed on to their o##spring.

The #orces o# natural selection act on phenot&pes" but onl& i# there is a change in the genot&pes o# a population has e$olution occurred. Population genetics is the stud& o# the genot&pes and their changes in entire populations. ,ink to a discussion.

The Measure of " itness"

!itness is a measure o# reproductive success. Those indi$iduals who lea$e the largest number o# mature o##spring are the #ittest. This can be achie$ed in se$eral wa&s Survival or mortality selection .ating success or sexual selection !amil& si4e or fecundity selection

Survival
0n& trait that promotes sur$i$al ' at least until one7s reproducti$e &ears are o$er ' increases #itness. (uch traits are adaptations.

Sexual Selection
In se)ual selection" one se) ' usuall& the #emale ' chooses among the a$ailable males. 0n& inherited trait that impro$es the mating success o# certain indi$iduals will become more pronounced in succeeding generations. (ome e)amples 8hen read& to mate" #emale three9spined sticklebacks 5#ish6 choose males with man& Class II .%C alleles o$er males with #ewer alleles. Class II alleles encode the proteins that present antigens to the immune s&stem. 3resumabl&" the more o# them &ou ha$e" the greater the di$ersit& o# parasite antigens &our immune s&stem can recogni4e and de#end against. ,ink to discussion o# how pol&morphism o# .%C alleles protects against parasites. The #emales distinguish between the males b& soluble molecules 5/odors/6 the males release into the water. %ow these /odors/ are controlled b& the .%C alleles is not known.

0 culture o# Drosophila set up with e:ual numbers o# red9e&ed and white9e&ed #lies o# both se)es will" a#ter ;5 generations or so" end up ha$ing onl& red9e&ed 5the /normal/6 #lies in it. This" despite the #act that white9e&ed #lies are just as health& and li$e just as long as red9e&ed #lies" i.e." the& are e:ual in terms o# sur$i$al. <ut" as it turns out" not onl& do red9e&ed #emales pre#er red9e&ed males" but white9e&ed #emales do also.

In other cases o# se)ual selection" one phenot&pe pre#ers to mate with others o# the same phenot&pe. This is called assortative mating.

ecundity Selection
The production o# a large number o# mature o##spring is a measure o# #itness. I stress mature because onl& the& can pass these traits on to another generation. (ome wa&s to do this Earlier breeding. I# some #emales become se)uall& mature earlier than others" their chances o# lea$ing o##spring are enhanced. !or some species 5e.g." #ish" o&sters6" which pro$ide little or no care #or their &oung" #itness is measured b& the number o# #ertili4ed eggs the& produce. !or species 5such as oursel$es6 that take care o# their &oung+ selection acts to reduce #amil& si4e 5to a point6. 0 large stud& in =tah 5=.(.06 showed that in the 19th centur& #amilies with #ewer children had more sur$i$ing grandchildren.

Are !umans Exempt from Natural Selection"

It has been argued that ad$ances in medicine" sanitation" etc. ha$e remo$ed humans #rom the rigors o# natural selection. There is probabl& some truth to this" but consider o# all the human eggs that are #ertili4ed" #ewer than hal# will e$er reproduce themsel$es. The others are eliminated as #ollows Mortality selection 0ppro)imatel& >?@ o# pregnancies end b& spontaneous abortion o# embr&os and #etuses or b& stillbirth. o Death in in#anc& and childhood claims another 5@ or more. Sexual selection o 0nother ;?@ will sur$i$e to adulthood but ne$er marr&. ecundity selection o *# those that do marr&" 1?@ will ha$e no children.
o

#ontinuous $ariation

.ost traits in a population. e.g." height bod& weight $ar& in a continuous wa& #rom one e)treme to the other. 0 plot o# the distribution o# the trait in a population o#ten produces a bell9shaped cur$e like this one that shows the distribution o# heights among a group o# male secondar&9school seniors. (uch a distribution could arise #rom en$ironmental #actors ' perhaps the continuous height $ariation in the bo&s is simpl& a result o# $ariation in their diet as the& grew up. or

genetic #actors A2iewB ' tall parents tend to ha$e tall children

or ' most likel& ' both.

!erita%ility
*ne can sort out the relati$e contribution o# genetic and en$ironmental #actors b& comparing the range o# a trait in the o##spring compared with the a$erage $alue o# that trait in their parents. I# the o##spring o# selected parents occup& the same range as the entire population" en$ironmental #actors are working alone. The trait has a &ero herita%ility. E)ample The length o# the seeds o# a pure strain o# beans ma& $ar& o$er se$eral millimeters. %owe$er" i# e)tra9large beans are mated" the new crop shows no shi#t to a larger si4e. (o the heritabilit& o# length is 4ero. *n the other hand" i# the o##spring o# two e)tra9large mice are just as large as the& are" genes are probabl& at work. The trait is said to ha$e a heritabilit& o# 1.

The Effects of Selection on Populations

The pressures o# natural selection can a##ect the distribution o# phenot&pes in a population in se$eral wa&s.

Sta%ili&ing Selection
-atural selection o#ten works to weed out indi$iduals at both e)tremes o# a range o# phenot&pes resulting in the reproducti$e success o# those near the mean. In such cases" the result is to maintain the status 'uo. It is not alwa&s eas& to see wh& both e)tremes should be handicapped+ perhaps se)ual selection or liabilit& to predation is at work. In an& case" stabili4ing selection is common. In humans" #or e)ample" the incidence o# in#ant mortalit& is higher #or $er& hea$& as well as #or $er& light babies.

(irectional Selection
0 population ma& #ind itsel# in circumstances where indi$iduals occup&ing one e)treme in the range o# phenot&pes are #a$ored o$er the others. (ince 19C>" 3eter and Dosemar& Erant ' aided b& a succession o# colleagues ' ha$e studied Darwin7s #inches in the Ealapagos Islands. ,ink to drawings o# Darwin7s #inches. 8hen rain#all" and thus #ood" are plenti#ul" the ground #inches tend to

ha$e a $aried diet" e.g." eat seeds o# a range o# si4es+ show considerable $ariation in bod& and beak si4e 5large beaks are better #or large seeds but can handle small seeds as well as small beaks6.

!rom 19CF through 19CC" a se$ere drought struck the islands" with $irtuall& no rain#all #or o$er a &ear. This caused a precipitous decline in the production o# the seeds that are the dietar& mainsta& o# )eospi&a fortis" the medium ground #inch. The graph 5#rom 3. T. <oag and 3. D. Erant in Science *+, 8;" 19816 shows how the population declined #rom 1G?? to ;?? on the island o# Daphne .ajor" a tin& 51?9acre6 member o# the Ealapagos Islands. ,ink to map. *ne o# the plants to make it through the drought produces seeds in large" tough #ruits that are $irtuall& impossible #or birds with a beak smaller than 1?.5 mm to eat. (ampling the birds that died as well as those that sur$i$ed showed that the larger birds were #a$ored o$er the smaller ones+ those with larger beaks were #a$ored o$er those with smaller ones.

<eak length 5mm6 Dead birds 1?.F8

<eak depth 5mm6 9.G;

(ur$i$ors 11.?C 9.9F %ere" then was natural selection at work. <ut did it produce e$olution1 The answer turned out to be &es. 0s the population o# E. #ortis reco$ered a#ter the rains returned" the a$erage bod& si4e and beak depth o# their o##spring was greater than be#ore 5an increase o# GH5@ #or beak depth6. The bell9shaped cur$e had been shi#ted to the right ' directional selection. .ore recentl&" the Erants and colleagues at %ar$ard .edical (chool ha$e shown that beak -idth and depth in the ground #inches are correlated with the timing and intensit& o# e)pression o# the gene" Bmp4" 5that encodes bone morphogenetic protein9G6 in the tissue that will #orm the upper beak. Bmp4 e)pression appears earlier in de$elopment and with greater intensit& in the large9beaked Eeospi4a magnirostris 5the large ground #inch6 than in its smaller9beaked relati$es" Eeospi4a #ortis 5the medium ground #inch6 and Eeospi4a #uliginosa 5the small ground #inch6. (ee 0b4hano$" 0." et al." Science" > (eptember ;??G.

%owe$er" beak length is correlated with the intensit& o# e)pression o# the gene CaM that encodes the Ca;I9binding protein calmodulin in the tissue that will #orm the upper beak. CaM e)pression is much higher in the embr&onic tissue o# the cactus #inches 5E. scandens and E. conirostris ' both with long beaks ' 2iew6 than in their short9beaked relati$es" the ground #inches E. #ortis and E. magnirostris A2iewB. 5(ee 0b4hano$" 0." et al." Nature" > 0ugust ;??F.6

Industrial Melanism .an& species o# moths in the <ritish Isles began to become darker in color in the 19th centur&. The best9 studied e)ample is the peppered moth" .iston

%etularia. The moth gets its name #rom the scattered dark markings on its wings and bod&. In 18G9" a coal9black mutant was #ound near .anchester" England. 8ithin a centur&" this black #orm had increased to 9?@ o# the population in this region. The moth #lies at night and rests b& da& on tree trunks. In areas #ar #rom industrial acti$it&" the trunks o# trees are encrusted with lichens. 0s the photos show" the light #orm 5circled in red6 is practicall& in$isible against this background. In areas where air pollution is se$ere" the combination o# to)ic gases and soot has killed the lichens and blackened the trunks. 0gainst such a background" the light #orm stands out sharpl&. The moth is pre&ed upon b& birds that pluck it #rom its resting place b& da&. In polluted woods" the dark #orm has a much better chance o# sur$i$ing undetected. 8hen the English geneticist %. <. D. Jettlewell 5who supplied the photos6 released moths o# both t&pes in the woods" he obser$ed that birds did" indeed" eat a much higher #raction o# the light moths he released than o# the dark. (ince pollution abatement programs were put in place a#ter 8orld 8ar II" the light #orm has been making a comeback in the ,i$erpool and .anchester areas.

(isruptive Selection
In some circumstances" indi$iduals at both e)tremes o# a range o# phenot&pes are #a$ored o$er those in the middle. This is called disrupti$e selection. 0n e)ample The residues 5/tailings/6 o# mines o#ten contain such high concentrations o# to)ic metals 5e.g." copper" lead6 that most plants are unable to grow on them. %owe$er" some hard& species 5e.g. certain grasses6 are able to spread #rom the surrounding uncontaminated soil onto such waste heaps. These plants de$elop resistance to the to)ic metals while their abilit& to grow on uncontaminated soil decreases. <ecause grasses are wind pollinated" breeding between the resistant and nonresistant populations goes on. <ut e$identl&" disrupti$e selection is at work. %igher death rates o# both less resistant plants growing on contaminated soil and more resistant plants growing on uncontaminated soil leads to increasing di$ergence o# the populations into two subpopulations with the e)treme mani#estations o# this trait. The e$olutionar& signi#icance o# disrupti$e selection lies in the possibilit& that the gene pool ma& become split into two distinct gene pools. This ma& be a wa& in which nespecies are #ormed.

The #ormation o# one or more species #rom a single precursor species is called speciation. It is the topic o# a separate page. ,ink to (peciation. 8elcomeK-e)t (earch ;1 December ;??9

Adaptation
!rom 8ikipedia" the #ree enc&clopedia

Lump to na$igation" search This article is about the evolutionary process. For other uses, see Adaptation (disambiguation).
3art o# the <iolog& series on

Evolution

Mechanisms and processes Adaptation Eenetic dri#t Eene #low .utation -atural selection (peciation /esearch and history Introduction E$idence E$olutionar& histor& o# li#e %istor& ,e$el o# support .odern s&nthesis *bjections M Contro$ers& (ocial e##ect Theor& and #act Evolutionary %iology fields

Cladistics Ecological genetics E$olutionar& de$elopment E$olutionar& ps&cholog& .olecular e$olution 3h&logenetics 3opulation genetics (&stematics .iology portal N $ O d O e

Adaptation is the e$olutionar& process whereb& a population becomes better suited to its habitat.A1BA;B This process takes place o$er man& generations"A>B and is one o# the basic phenomena o# biolog&.AGB The term adaptation ma& also re#er to a #eature which is especiall& important #or an organism7s sur$i$al.A5B !or e)ample" the adaptation o# horses7 teeth to the grinding o# grass" or their abilit& to run #ast and escape predators. (uch adaptations are produced in a $ariable population b& the better suited #orms reproducing more success#ull&" that is" b& natural selection.

#ontents
AhideB

1 Eeneral principles o 1.1 De#initions o 1.; 0daptedness and #itness ; <rie# histor& > T&pes o# adaptation o >.1 Changes in habitat >.1.1 %abitat tracking >.1.; Eenetic change o >.; Intimate relationships co9adaptations >.;.1 .imicr& o >.> The basic machiner& internal adaptations o >.G Compromise and con#lict between adaptations G (hi#ts in #unction o G.1 3re9adaptations o G.; Co9option o# e)isting traits e)aptation 5 Delated issues o 5.1 -on9adapti$e traits 5.1.1 !itness landscapes+ dri#t 5.1.; 2estigial organs o 5.; E)tinction 5.;.1 Co9e)tinction

5.> !le)ibilit&" acclimati4ation" learning 5.>.1 !le)ibilit& 5.>.; 0cclimati4ation 5.>.> ,earning F !unction and teleonom& o F.1 !unction o F.; Teleonom& C (ee also
o

8 De#erences

0edit1 )eneral principles

The signi#icance o# an adaptation can onl& be understood in relation to the total biolog& o# the species. ulian !u"leyAFB 0daptation is" #irst o# all" a process" rather than a ph&sical part o# a bod&.ACB The distinction ma& be seen in an internal parasite 5such as a #luke6" where the bodil& structure is greatl& simpli#ied" but ne$ertheless the organism is highl& adapted to its unusual en$ironment. !rom this we see that adaptation is not just a matter o# $isible traits in such parasites critical adaptations take place in the li#e9c&cle" which is o#ten :uite comple).A8B %owe$er" as a practical term" adaptation is o#ten used #or the product those #eatures o# a species which result #rom the process. .an& aspects o# an animal or plant can be correctl& called adaptations" though there are alwa&s some #eatures whose #unction is in doubt. <& using the term adaptation #or the e$olutionar& process" and adaptive trait #or the bodil& part or #unction 5the product6" the two senses o# the word ma& be distinguished. 0daptation ma& be seen as one aspect o# a two9stage process. !irst" there is speciation 5species9splitting or cladogenesis6" caused b& geographical isolation or some other mechanism.A9BA1?B (econd" there #ollows adaptation" dri$en b& natural selection. (omething like this must ha$e happened with Darwin7s #inches" and there are man& other e)amples. The present #a$ourite is the e$olution o# cichlid #ish in 0#rican lakes" where the :uestion o# reproducti$e isolation is much more comple).A11BA1;B 0nother great principle is that an organism must be $iable at all stages o# its de$elopment and at all stages o# its e$olution. This is ob$iousl& true" and it #ollows that there are constraints on the e$olution o# de$elopment" beha$iour and structure o# organisms. The main constraint" o$er which there has been much debate" is the re:uirement that changes in the s&stem during e$olution should be relati$el& small changes" because the bod& s&stems are so comple) and interlinked. This is a sound principle" though there ma& be rare e)ceptions pol&ploid& in plants is common"A1>B and the s&mbiosis o# micro9 organisms that #ormed the eukar&ota is a more e)otic e)ample.A1GB 0ll adaptations help organisms sur$i$e in their ecological niches.A15B These adaptati$e traits ma& be structural" beha$ioral or ph&siological. (tructural adaptations are ph&sical

#eatures o# an organism 5shape" bod& co$ering" armament+ and also the internal organi4ation6. <eha$ioural adaptations are composed o# inherited beha$iour chains andMor the abilit& to learn beha$iours ma& be inherited in detail 5instincts6" or a tendenc& #or learning ma& be inherited 5see neurops&cholog&6. E)amples searching #or #ood" mating" $ocali4ations. 3h&siological adaptations permit the organism to per#orm special #unctions 5#or instance" making $enom" secreting slime" phototropism6+ but also more general #unctions such as growth and de$elopment" temperature regulation" ionic balance and other aspects o# homeostasis. 0daptation" then" a##ects all aspects o# the li#e o# an organism.

0edit1 (efinitions
The #ollowing de#initions are mainl& due to Theodosius Dob4hansk&. 1. Adaptation is the e$olutionar& process whereb& an organism becomes better able to li$e in its habitat or habitats.A1FB ;. Adaptedness is the state o# being adapted the degree to which an organism is able to li$e and reproduce in a gi$en set o# habitats.A1CB >. 0n adaptive trait is an aspect o# the de$elopmental pattern o# the organism which enables or enhances the probabilit& o# that organism sur$i$ing and reproducing.A18B

0edit1 Adaptedness and fitness

.ain article !itness 5biolog&6 !rom the abo$e de#initions" it is clear that there is a relationship between adaptedness and #itness 5a ke& population genetics concept6. !itness is an estimate and a predictor o# the rate o# natural selection. 8hat natural selection does is change the relati$e #re:uencies o# alternati$e phenot&pes" inso#ar as the& are heritable.A19B 0lthough the two are connected" the one does not impl& the other a phenot&pe with high adaptedness ma& not ha$e high #itness. Dob4hansk& mentioned the e)ample o# the Cali#ornian redwood" which is highl& adapted" but a relic species in danger o# e)tinction.A1FB Elliott (ober commented that adaptation was a retrospecti$e concept since it implied something about the histor& o# a trait" whereas #itness predicts a trait7s #uture.A;?B 1. !itness. The degree o# demographic di##erence among phenot&pes. =suall& a relati$e measure the a$erage contribution to a breeding population b& a phenot&pe or a class o# phenot&pes. This is also known as $ar%inian #itness" relative #itness" selective coe##icient" and other terms. ;. 0daptedness. =suall& an absolute measure the a$erage absolute contribution to the breeding population b& a carrier o# a phenot&pe or a class o# phenot&pes. 0lso known as absolute #itness" and as the .althusian parameter when applied to species as a whole.A;1B

0edit1 .rief history

.ain article %istor& o# e$olutionar& thought 0daptation as a #act o# li#e has been accepted b& all the great thinkers who ha$e tackled the world o# li$ing organisms. It is their e)planations o# how adaptation arises that separates these thinkers. 0 #ew o# the most signi#icant ideas A;;B

Empedocles did not belie$e that adaptation re:uired a #inal cause 5P purpose6" but /came about naturall&" since such things sur$i$ed/. 0ristotle" howe$er" did belie$e in #inal causes. In natural theolog&" adaptation was interpreted as the work o# a deit&" e$en as e$idence #or the e)istence o# Eod.A;>B 8illiam 3ale& belie$ed that organisms were per#ectl& adapted to the li$es the& lead" an argument that shadowed ,eibni4" who had argued that Eod had brought about the best o# all possible worlds. 2oltaire7s Dr 3anglossA;GB is a parod& o# this optimistic idea" and %ume also argued against design.A;5B The Bridge%ater Treatises are a product o# natural theolog&" though some o# the authors managed to present their work in a #airl& neutral manner. The series was lampooned b& Dobert Jno)" who held :uasi9e$olutionar& $iews" as the Bilge%ater Treatises. Darwin broke with the tradition b& emphasising the #laws and limitations which occurred in the animal and plant worlds.A;FB

,amarck ,amarck. %is is a proto9e$olutionar& theor& o# the inheritance o# ac:uired traits" whose main purpose is to e)plain adaptations b& natural means.A;CB %e proposed a tendenc& #or organisms to become more comple)" mo$ing up a ladder o# progress" plus /the in#luence o# circumstances/" usuall& e)pressed as use and disuse. %is e$olutionar& ideas" and those o# Eeo##ro&" #ail because the& cannot be reconciled with heredit&. This was known e$en be#ore .endel b& medical men interested in human races 58ells" ,awrence6" and especiall& b& 8eismann. .an& other students o# natural histor&" such as <u##on" accepted adaptation" and some also accepted e$olution" without $oicing their opinions as to the mechanism. This illustrates the real merit o# Darwin and 8allace" and secondar& #igures such as <ates" #or putting #orward a mechanism whose signi#icance had onl& been glimpsed pre$iousl&. 0 centur& later" e)perimental #ield studies and breeding e)periments b& such as !ord and Dob4hansk& produced e$idence that natural selection was not onl& the 7engine7 behind adaptation" but was a much stronger #orce than had pre$iousl& been thought.A;8BA;9BA>?B

0edit1 Types of adaptation

0daptation is the heart and soul o# e$olution. &iles 'ldredgeA>1B

0edit1 #hanges in ha%itat

<e#ore Darwin" adaptation was seen as a #i)ed relationship between an organism and its habitat. It was not appreciated that as the climate changed" so did the habitat+ and as the habitat changed" so did the biota. 0lso" habitats are subject to changes in their biota #or e)ample" in$asions o# species #rom other areas. The relati$e numbers o# species in a gi$en habitat are alwa&s changing. Change is the rule" though much depends on the speed and degree o# the change. 8hen the habitat changes" three main things ma& happen to a resident population habitat tracking" genetic change or e)tinction. In #act" all three things ma& occur in se:uence. (# these three e##ects, only genetic change brings about adaptation. 0edit1 !a%itat trac2ing 8hen a habitat changes" the most common thing to happen is that the resident population mo$es to another locale which suits it+ this is the t&pical response o# #l&ing insects or oceanic organisms" who ha$e wide 5though not unlimited6 opportunit& #or mo$ement.A>;B This common response is called habitat trac)ing. It is one e)planation put #orward #or the periods o# apparent stasis in the #ossil record 5the punctuated e:uilibrium thesis6.A>>B 0edit1 )enetic change Eenetic change is what occurs in a population when natural selection acts on the genetic $ariabilit& o# the population. <& this means" the population adapts geneticall& to its circumstances.A>GB Eenetic changes ma& result in $isible structures" or ma& adjust ph&siological acti$it& in a wa& that suits the changed habitat. It is now clear that habitats and biota do #re:uentl& change. There#ore" it #ollows that the process o# adaptation is ne$er #inall& complete.A>5B *$er time" it ma& happen that the en$ironment changes little" and the species comes to #it its surroundings better and better. *n the other hand" it ma& happen that changes in the en$ironment occur relati$el& rapidl&" and then the species becomes less and less well adapted. (een like this" adaptation is a genetic trac)ing process" which goes on all the time to some e)tent" but especiall& when the population cannot or does not mo$e to another" less hostile area. 0lso" to a greater or lesser e)tent" the process a##ects e$er& species in a particular ecos&stem.A>FBA>CB 2an 2alen thought that e$en in a stable en$ironment" competing species had to constantl& adapt to maintain their relati$e standing. This became known as the Ded Queen h&pothesis.

0edit1 Intimate relationships3 co4adaptations

.ain article Co9adaptation In co9e$olution" where the e)istence o# one species is tightl& bound up with the li#e o# another species" new or 7impro$ed7 adaptations which occur in one species are o#ten #ollowed b& the appearance and spread o# corresponding #eatures in the other species. There are man& e)amples o# this+ the idea emphasises that the li#e and death o# li$ing things is intimatel& connected" not just with the ph&sical en$ironment" but with the li#e o# other species. These relationships are intrinsicall& d&namic" and ma& continue on a trajector& #or millions o# &ears" as has the relationship between #lowering plants and insects 5pollination6. 3ollinator constanc& these hone&bees selecti$el& $isit #lowers #rom onl& one species" as can be seen b& the colour o# the pollen in their baskets

Co9e)tinction In#ection9resistance .imicr& .utualism 3arasite9host 3ollination s&ndrome 3redator9pre& (&mbiosis

The gut contents" wing structures" and mouthpart morphologies o# #ossili4ed beetles and #lies suggest that the& acted as earl& pollinators. The association between beetles and angiosperms during the earl& Cretaceous period led to parallel radiations o# angiosperms and insects into the late Cretaceous. The e$olution o# nectaries in late Cretaceous #lowers signals the beginning o# the mutualism between h&menopterans and angiosperms.A>8B 0edit1 Mimicry .ain article .imicr&

0 and < show real wasps+ the rest are mimics three ho$er#lies and one beetle. %enr& 8alter <ates7 work on 0ma4onian butter#lies led him to de$elop the #irst scienti#ic account o# mimicr&" especiall& the kind o# mimicr& which bears his name <atesian mimicr&.A>9B This is the mimicr& b& a palatable species o# an unpalatable or no)ious species. 0 common e)ample seen in temperate gardens is the ho$er9#l&" man& o# which H though bearing no sting H mimic the warning colouration o# h&menoptera 5wasps and bees6. (uch mimicr& does not need to be per#ect to impro$e the sur$i$al o# the palatable species.AG?B <ates" 8allace and .Rller belie$ed that <atesian and .Rllerian mimicr& pro$ided e$idence #or the action o# natural selection" a $iew which is now standard amongst biologists.AG1B 0ll aspects o# this situation can be" and ha$e been" the subject o# research.AG;B !ield and e)perimental work on these ideas continues to this da&+ the topic connects strongl& to speciation" genetics and de$elopment.AG>B

.ore on mimicr& 8arning Colour and .imicr& ,ecture outline #rom =ni$ersit& College ,ondon

0edit1 The %asic machinery3 internal adaptations

There are some important adaptations to do with the o$erall coordination o# the s&stems in the bod&. (uch adaptations ma& ha$e signi#icant conse:uences. E)amples" in $ertebrates" would be temperature regulation" or impro$ements in brain #unction" or an e##ecti$e immune s&stem. 0n e)ample in plants would be the de$elopment o# the reproducti$e s&stem in #lowering plants.AGGB (uch adaptations ma& make the clade 5monoph&letic group6 more $iable in a wide range o# habitats. The ac:uisition o# such major adaptations has o#ten ser$ed as the spark #or adapti$e radiation" and huge success #or long periods o# time #or a whole group o# animals or plants.

0edit1 #ompromise and conflict %et-een adaptations

It is a pro#ound truth that -ature does not know best+ that genetical e$olution... is a stor& o# waste" makeshi#t" compromise and blunder. *eter Meda%arAG5B 0ll adaptations ha$e a downside horse legs are great #or running on grass" but the& can7t scratch their backs+ mammals7 hair helps temperature" but o##ers a niche #or ectoparasites+ the onl& #l&ing penguins do is under water. 0daptations ser$ing di##erent #unctions ma& be mutuall& destructi$e. Compromise and make9shi#t occur widel&" not per#ection. (election pressures pull in di##erent directions" and the adaptation that results is some kind o# compromise.AGFB (ince the phenot&pe as a whole is the target o# selection" it is impossible to impro$e simultaneousl& all aspects o# the phenot&pe to the same degree. 'rnst Mayr.AGCB Consider the antlers o# the Irish elk" 5o#ten supposed to be #ar too large+ in deer antler si4e has an allometric relationship to bod& si4e6. *b$iousl& antlers ser$e positi$el& #or de#ence against predators" and to score $ictories in the annual rut. <ut the& are costl& in terms o# resource. Their si4e during the last glacial period presumabl& depended on the relati$e gain and loss o# reproducti$e capacit& in the population o# elks during that time. AG8B 0nother e)ample camou#lage to a$oid detection is destro&ed when $i$id colors are displa&ed at mating time. %ere the risk to li#e is counterbalanced b& the necessit& #or reproduction.

0n Indian 3eacock7s train in #ull displa& The peacock7s ornamental train 5grown anew in time #or each mating season6 is a #amous adaptation. It must reduce his maneu$erabilit& and #light" and is hugel& conspicuous+ also" its growth costs #ood resources. Darwin7s e)planation o# its ad$antage was in terms o# se)ual selection /it depends on the ad$antage which certain indi$iduals ha$e o$er other indi$iduals o# the same se) and species" in e)clusi$e relation to reproduction./AG9B The kind o# se)ual selection represented b& the peacock is called 7mate choice7" with an implication that the process selects the more #it o$er the less #it" and so has sur$i$al $alue.A5?B The recognition o# se)ual selection was #or a long time in abe&ance" but has been rehabilitated.A51B In practice" the blue pea#owl *avo cristatus is a prett& success#ul species" with a big natural range in India" so the o$erall outcome o# their mating s&stem is :uite $iable.

The con#lict between the si4e o# the human #oetal brain at birth" 5which cannot be larger than about G??ccs" else it will not get through the mother7s pel$is6 and the si4e needed #or an adult brain 5about 1G??ccs6" means the brain o# a newborn child is :uite immature. The most $ital things in human li#e 5locomotion" speech6 just ha$e to wait while the brain grows and matures. That is the result o# the birth compromise. .uch o# the problem comes #rom our upright bipedal stance" without which our pel$is could be shaped more suitabl& #or birth. -eanderthals had a similar problem.A5;BA5>BA5GB

0edit1 Shifts in function

0daptation and #unction are two aspects o# one problem. ulian !u"leyA55B

0edit1 Pre4adaptations
This occurs when a species or population has characteristics which 5b& chance6 are suited #or conditions which ha$e not &et arisen. !or e)ample" the pol&ploid rice9grass +partina to%nsendii is better adapted than either o# its parent species to their own habitat o# saline marsh and mud9#lats.A5FB 8hite ,eghorn #owl are markedl& more resistant to $itamin < de#icienc& than other breeds.A5CB *n a plenti#ul diet there is no di##erence" but on a restricted diet this preadaptation could be decisi$e. 3re9adaptation ma& occur because a natural population carries a huge :uantit& o# genetic $ariabilit&.A58B In diploid eukar&otes" this is a conse:uence o# the s&stem o# se)ual reproduction" where mutant alleles get partiall& shielded" #or e)ample" b& the selecti$e ad$antage o# hetero4&gotes. .icro9organisms" with their huge populations" also carr& a great deal o# genetic $ariabilit&. The #irst e)perimental e$idence o# the pre9adapti$e nature o# genetic $ariants in micro9 organisms was pro$ided b& (al$ador ,uria and .a) DelbrRck who de$eloped #luctuation anal&sis" a method to show the random #luctuation o# pre9e)isting genetic changes that con#erred resistance to phage in the bacterium 'scherichia coli.

0edit1 #o4option of existing traits3 exaptation

.ain article E)aptation The classic e)ample is the ear ossicles o# mammals" which we know #rom palaeontological and embrological studies originated in the upper and lower jaws and the h&oid o# their (&napsid ancestors" and #urther back still were part o# the gill arches o# earl& #ish.A59BAF?B 8e owe this esoteric knowledge to the comparati$e anatomists" who" a centur& ago" were at the cutting edge o# e$olutionar& studies.AF1B The word e"aptation was coined to co$er these shi#ts in #unction" which are surprisingl& common in e$olutionar& histor&.AF;B The origin o# wings #rom #eathers that were originall& used #or temperature regulation is a more recent disco$er& 5see #eathered dinosaurs6.

0edit1 /elated issues

0edit1 Non4adaptive traits
(ome traits appear to be not adapti$e" that is" selecti$el& neutral. There ma& be $arious causes the utilit& o# a trait is lost and does not now appear adapti$e+ the utilit& o# a trait is unknown+ the trait is a conse:uence o# another trait that is adapti$e 5i.e. spandrels6. <ecause genes ha$e pleiotropic e##ects" not all traits ma& be #unctional. *# course" a trait ma& ha$e been adapti$e at some point in an organism7s e$olutionar& histor&" but habitats change" leading to adaptations becoming redundant or e$en a hindrance 5maladaptations6. (uch adaptations are termed $estigial. (o" the utilit& o# adaptations ma& ebb and #low. 0edit1 itness landscapes5 drift .ain article !itness landscape .ain article Eenetic dri#t (ewall 8right7s e)planation #or e$olutionar& stasis was that organisms come to occup& adaptive pea)s. In order to e$ol$e to another" higher peak" the species would #irst ha$e to pass through a $alle& o# maladapti$e intermediate stages. This could happen b& genetic dri#t i# the population were small enough. This was 8right7s shi#ting balance theory o# evolution.AF>B There has been much skepticism among e$olutionar& biologists as to whether these rather delicate conditions hold o#ten in natural populations.A58B Donald !isher #elt that most populations in nature were too large #or these e##ects o# genetic dri#t to be important.A;8B 0edit1 $estigial organs .ain article 2estigialit& .an& organisms ha$e $estigial organs" which are the remnants o# #ull& #unctional structures in their ancestors. 0s a result o# changes in li#est&le the organs became redundant" and are either not #unctional or reduced in #unctionalit&. 8ith the loss o# #unction goes the loss o# positi$e selection" and the subse:uent accumulation o# deleterious mutations. (ince an& structure represents some kind o# cost to the general econom& o# the bod&" an ad$antage ma& accrue #rom their elimination once the& are not #unctional. E)amples wisdom teeth in humans+ the loss o# pigment and #unctional e&es in ca$e #auna+ the loss o# structure in endoparasites.AFGB

0edit1 Extinction
.ain article E)tinction I# a population cannot mo$e or change su##icientl& to preser$e its long9term $iabilit&" then ob$iousl&" it will become e)tinct" at least in that locale. The species ma& or ma& not sur$i$e in other locales. (pecies e)tinction occurs when the death rate o$er the entire

species 5population" gene pool ...6 e)ceeds the birth rate #or a long enough period #or the species to disappear. It was an obser$ation o# 2an 2alen that groups o# species tend to ha$e a characteristic and #airl& regular rate o# e)tinction.AF5B 0edit1 #o4extinction .ain article Co9e)tinction Lust as we ha$e co9adaptation" there is also co9e)tinction. Co9e)tinction re#ers to the loss o# a species due to the e)tinction o# another+ #or e)ample" the e)tinction o# parasitic insects #ollowing the loss o# their hosts. Co9e)tinction can also occur when a #lowering plant loses its pollinator" or through the disruption o# a #ood chain.AFFB /(pecies co9 e)tinction is a mani#estation o# the interconnectedness o# organisms in comple) ecos&stems ... 8hile co9e)tinction ma& not be the most important cause o# species e)tinctions" it is certainl& an insidious one/.AFCB

0edit1 lexi%ility6 acclimati&ation6 learning

Fle"ibility deals with the relati$e capacit& o# an organism to maintain themsel$es in di##erent habitats their degree o# speciali4ation. Acclimati,ation is a term used #or automatic ph&siological adjustments during li#e+ learning is the term used #or impro$ement in beha$ioral per#ormance during li#e. In biolog& these terms are pre#erred" not adaptation" #or changes during li#e which impro$e the per#ormance o# indi$iduals. These adjustments are not inherited geneticall& b& the ne)t generation. 0daptation" on the other hand" occurs o$er man& generations+ it is a gradual process caused b& natural selection which changes the genetic make9up o# a population so the collecti$e per#orms better in its niche. 0edit1 lexi%ility 3opulations di##er in their phenot&pic plasticit&" which is the abilit& o# an organism with a gi$en genot&pe to change its phenot&pe in response to changes in its habitat" or to its mo$e to a di##erent habitat.AF8BAF9B To a greater or lesser e)tent" all li$ing things can adjust to circumstances. The degree o# #le)ibilit& is inherited" and $aries to some e)tent between indi$iduals. 0 highl& speciali4ed animal or plant li$es onl& in a well9de#ined habitat" eats a speci#ic t&pe o# #ood" and cannot sur$i$e i# its needs are not met. .an& herbi$ores are like this+ e)treme e)amples are koalas which depend on eucal&ptus" and pandas which re:uire bamboo. 0 generalist" on the other hand" eats a range o# #ood" and can sur$i$e in man& di##erent conditions. E)amples are humans" rats" crabs and man& carni$ores. The tendency to beha$e in a speciali4ed or e)plorator& manner is inherited H it is an adaptation. Dather di##erent is developmental #le"ibility- /0n animal or plant is de$elopmentall& #le)ible i# when it is raised or trans#erred to new conditions it de$elops so that it is better

#itted to sur$i$e in the new circumstances/.AC?B *nce again" there are huge di##erences between species" and the capacities to be #le)ible are inherited. 0edit1 Acclimati&ation .ain article 0cclimati4ation I# humans mo$e to a higher altitude" respiration and ph&sical e)ertion become a problem" but a#ter spending time in high altitude conditions the& acclimati,e to the pressure b& increasing production o# red blood corpuscles. The ability to acclimati4e is an adaptation" but not the acclimati4ation itsel#. !ecundit& goes down" but deaths #rom some tropical diseases also goes down. *$er a longer period o# time" some people will reproduce better at these high altitudes than others. The& will contribute more hea$il& to later generations. Eraduall& the whole population becomes adapted to the new conditions. This we know takes place" because the per#ormance o# long9term communities at higher altitude is signi#icantl& better than the per#ormance o# new arri$als" e$en when the new arri$als ha$e had time to make ph&siological adjustments.AC1B (ome kinds o# acclimati4ation happen so rapidl& that the& are better called re#le)es. The rapid colour changes in some #lat#ish" cephalopods" chameleons are e)amples.AC;B 0edit1 7earning (ocial learning is supreme #or humans" and is possible #or :uite a #ew mammals and birds o# course" that does not in$ol$e genetic transmission e)cept to the e)tent that the capacities are inherited. (imilarl&" the capacity to learn is an inherited adaptation" but not what is learnt+ the capacit& #or human speech is inherited" but not the details o# language.

0edit1 unction and teleonomy

0daptation raises some issues concerning how biologists use ke& terms such as #unction.

0edit1 unction
To sa& something has a #unction is to sa& something about what it does #or the organism" ob$iousl&. It also sa&s something about its histor& how it has come about. 0 heart pumps blood that is its #unction. It also emits sound" which is just an ancillar& side9e##ect. That is not its #unction. The heart has a histor& 5which ma& be well or poorl& understood6" and that histor& is about how natural selection #ormed and maintained the heart as a pump. E$er& aspect o# an organism that has a #unction has a histor&. -ow" an adaptation must ha$e a #unctional histor& there#ore we e)pect it must ha$e undergone selection caused b& relati$e sur$i$al in its habitat. It would be :uite wrong to use the word adaptation about a trait which arose as a b&9product.AC>BACGB

It is widel& regarded as unpro#essional #or a biologist to sa& something like /0 wing is #or #l&ing/" although that is their normal #unction. 0 biologist would be conscious that sometime in the remote past #eathers on a small dinosaur had the #unction o# retaining heat" and that later man& wings were not used #or #l&ing 5e.g. penguins" ostriches6. (o" the biologist would rather sa& that the wings on a bird or an insect usuall& had the #unction o# aiding #light. That would carr& the connotation o# being an adaptation with a histor& o# e$olution b& natural selection.

0edit1 Teleonomy
.ain article Teleonom& Teleonom& is a term in$ented to describe the stud& o# goal9directed #unctions which are not guided b& the conscious #orethought o# man or an& supernatural entit&. It is contrasted with 0ristotle7s teleolog&" which has connotations o# intention" purpose and #oresight. E$olution is teleonomic+ adaptation hoards hindsight rather than #oresight. The #ollowing is a de#inition #or its use in biolog& Teleonom& The h&pothesis that adaptations arise without the e)istence o# a prior purpose" but b& the action o# natural selection on genetic $ariabilit&.AC5B The term ma& ha$e been suggested b& Colin 3ittendrigh in 1958+ACFB it grew out o# c&bernetics and sel#9organising s&stems. Ernst .a&r" Eeorge C. 8illiams and Lac:ues .onod picked up the term and used it in e$olutionar& biolog&.ACCBAC8BAC9BA8?B 3hilosophers o# science ha$e also commented on the term. Ernest -agel anal&sed the concept o# goal9directedness in biolog&+A81B and Da$id %ull commented on the use o# teleolog& and teleonom& b& biologists %aldane can be #ound remarking" /Teleolog& is like a mistress to a biologist he cannot li$e without her but heSs unwilling to be seen with her in public/. Toda& the mistress has become a law#ull& wedded wi#e. <iologists no longer #eel obligated to apologi4e #or their use o# teleological language+ the& #launt it. The onl& concession which the& make to its disreputable past is to rename it Tteleonom&S.A8;B

0edit1 See also

Evolutionary biology portal

0dapti$e radiation Co9adaptation Co9e$olution Ecological trap E)aptation Intragenomic con#lict

.aladaptation .imicr& 3ol&morphism 5biolog&6

0edit1 /eferences
1. 8 The ("#ord $ictionary o# +cience de#ines adaptation as /0n& change in the structure or #unctioning o# an organism that makes it better suited to its en$ironment/. ;. 8 <owler 3.L. ;??>. 'volution- the history o# an idea. Cali#ornia. p1? >. 8 3atterson C. 1999. 'volution. -atural %istor& .useum" ,ondon. p1 G. 8 8illiams" Eeorge C. 19FF. Adaptation and natural selection- a criti.ue o# some current evolutionary thought. 3rinceton. /E$olutionar& adaptation is a phenomenon o# per$asi$e importance in biolog&./ p5 5. 8 <oth uses o# the term 7adaptation7 are recogni4ed b& Jing D.C. (tans#ield 8.D. and .ulligan 3. ;??F. A dictionary o# genetics. *)#ord" Cth ed. F. 8 %u)le&" Lulian 19G;. 'volution the modern synthesis. 0llen K =nwin" ,ondon. pGG9 C. 8 .a&r" Ernst 198;. The gro%th o# biological thought. %ar$ard. pG8> /0daptation... could no longer be considered a static condition" a product o# a creati$e past" and became instead a continuing d&namic process./ 8. 8 3rice 3.8. 198?. The evolutionary biology o# parasites. 3rinceton. 9. 8 .a&r E. 19F>. Animal species and evolution. %ar$ard. 1?. 8 .a&r" Ernst 198;. The gro%th o# biological thought- diversity, evolution and inheritance. %ar$ard. p5F;H5FF 11. 8 (al4burger 8." .ack T." 2erhe&en E." .e&er 0. 5;??56. /*ut o# Tangan&ika Eenesis" e)plosi$e speciation" ke&9inno$ations and ph&logeograph& o# the haplochromine cichlid #ishes/ 53D!6. BMC 'volutionary Biology 9 516 1C. doi 1?.118FM1GC19;1G89591C. 3.ID 15C;>F98. 3.C 55GCCC. http MMwww.biomedcentral.comMcontentMpd#M1GC19;1G89591C.pd#. 1;. 8 Jorn#ield" Ir$+ (mith" 3eter 5-o$ember ;???6. /0#rican Cichlid !ishes .odel (&stems #or E$olutionar& <iolog&/. Annual /evie% o# 'cology and +ystematics :+ 1F>. doi 1?.11GFMannure$.ecols&s.>1.1.1F>. http MMarjournals.annualre$iews.orgMdoiMabsM1?.11GFMannure$.ecols&s.>1.1.1F>. 1>. 8 (tebbins" E. ,ed&ard" Lr. 195?. 0ariation and evolution in plants. Columbia. *olyploidy" chapters 8 and 9. 1G. 8 .argulis" ,&nn 5ed6 1991. +ymbiosis as a source o# evolutionary innovationspeciation and morphogenesis .IT. I(<- ?9;F;91>;F999 15. 8 %utchinson E. E$el&n 19F5. The ecological theatre and the evolutionary play. Uale. The niche is the central concept in e$olutionar& ecolog&+ see especiall& part II The niche an abstractl& inhabited h&per$olume. p;FHC8 1F. V a b Dob4hansk& T. 19F8. *n some #undamental concepts o# e$olutionar& biolog&. 'volutionary biology *" 1H>G. 1C. 8 Dob4hansk& T. 19C?. 1enetics o# the evolutionary process. Columbia" -.U. pGH F" C9H8;" 8GH8C

18. 8 Dob4hansk& T. 195F. Eenetics o# natural populations WW2. Eenetic changes in populations o# $rosophila pseudoobscura and $rosphila persimilis in some locations in Cali#ornia. 'volution +;" 8;H9;. 19. 8 Endler" Lohn 0. 198F. &atural selection in the %ild. 3rinceton. p>>H51 7!itness and adaptation7. ;?. 8 (ober" Elliott 198G. The nature o# selection- a philosophical en.uiry. ..I.T. ;1. 8 #ollowing discussion in Endler" Lohn 0. 198F. &atural selection in the %ild. 3rinceton. p>>H51 7!itness and adaptation7. ;;. 8 re#erences and details in their articles ;>. 8 Desmond" 0drian 1989. The politics o# evolution. Chicago. p>1M>;" #ootnote 18. ;G. 8 In Candide, ou l2optimisme. ;5. 8 (ober" Elliott 199>. *hilosophy o# biology. *)#ord. Chapter ; ;F. 8 Darwin" Charles. 18C;. The origin o# species. Fth edition" p>9C Dudimentar&" atrophied and aborted organs. ;C. 8 see" #or e)ample" the discussion in <owler" 3eter %. ;??>. 'volution- the history o# an idea. >rd ed" Cali#ornia. p8FH95" especiall& /8hate$er the true nature o# ,amark7s theor&" it was his mechanism o# adaptation that caught the attention o# later naturalists/. 5p9?6 ;8. V a b 3ro$ine" 8illiam 198F. +e%all 3right and evolutionary biology. =ni$ersit& o# Chicago 3ress. ;9. 8 !ord E.<. 19C5. 'cological genetics" Gth ed. Chapman and %all" ,ondon. >?. 8 *rr %. ;??5. The genetic theor& o# adaptation a brie# histor&. &ature /ev. 1enetics <" ;" p119H1;C. >1. 8 Eldredge" -iles 1995. /einventing $ar%in- the great evolutionary debate. 8ile& -.U. p>> >;. 8 Eldredge" -iles 198F. Time #rames- the rethin)ing o# $ar%inian evolution and the theory o# punctuated e.uilibria. p1>F" (# glaciers and beetles. >>. 8 Eldredge" -iles 1995. /einventing $ar%in- the great evolutionary debate. 8ile&" -.U. pFG >G. 8 *rr %. ;??5. The genetic theor& o# adaptation a brie# histor&. &ature /ev. 1enetics" <" 119H1;C. >5. 8 .a&r" Ernst 198;. The gro%th o# biological thought- diversity, evolution and inheritance. !arvard. %ar$ard. pG81 5and se:uence6 tells how Darwin7s ideas on adaptation de$eloped as he came to appreciate it as /a continuing d&namic process/ 5bottom pG8>6. >F. 8 (tereln& J. K Eri##iths 3.E. 1999. +e" and death- an introduction to philosophy o# biology. =ni$ersit& o# Chicago 3ress. p;1C I(<- *9;;F9CC>?G9> >C. 8 !reeman (. K %erron L.C. ;??C. 'volutionary analysis. 3earson Education. p>FG I(<- ?91>9;;C58G98 >8. 8 (tebbins" E. ,ed&ard" Lr. 19CG. Flo%ering plants- evolution above the species level. %ar$ard. >9. 8 Carpenter ED% and !ord E< 19>>. Mimicry. .ethuen" ,ondon. G?. 8 8ickler 8. 19F8. Mimicry in plants and animals. 8orld =ni$ersit& ,ibrar&" ,ondon. G1. 8 .oon %.3. 19CF. !enry 3alter Bates F/+ 456784596- e"plorer, scientist and dar%inian. ,eicestershire .useums" ,eicester.

G;. 8 Du)ton ED" (herratt T- and (peed .3 ;??G. Avoiding attac)- the evolutionary ecology o# crypsis, %arning signals and mimicry. *)#ord. G>. 8 .allet" Lames ;??1. The speciation re$olution. 'volutionary Biology +," 88C9 8. GG. 8 (tebbins" E. ,ed&ard" Lr. 19CG. Flo%ering plants- evolution above the species level. %ar$ard. Contains an e)tensi$e anal&sis o# the e$olution o# adaptations in the radiation o# 0ngiosperms. G5. 8 .edawar" 3eter 19F?. The #uture o# Man. .ethuen" ,ondon. GF. 8 Lacob" !rancois 19CC. E$olution and tinkering. +cience +=< 11F1H11FF. GC. 8 .a&r" Ernst 198;. The gro%th o# biological thought- diversity, evolution and inheritance. %ar$ard. p589 G8. 8 It is" o# course" not possible to test selecti$e pressures on e)tinct populations in an& direct wa&. E*=,D" (TE3%E- L. 519CG6 *rigin and #unction o# 7bi4arre7 structures 9 antler si4e and skull si4e in 7Irish Elk7" Megaloceros giganteus. 'volution *>5;6 1919;;?. doi 1?.;>?CM;G?C>;; 5!irst page te)t6 G9. 8 Darwin" Charles 18C1. The $escent o# Man and selection in relation to se". .urra&" ,ondon. 5?. 8 The case was treated b& !isher D.0. 19>?. 1enetical theory o# natural selection. *)#ord. p1>GH1>9. 51. 8 Cronin" %elen 1991. The ant and the peacoc)- altruism and se"ual selection #rom $ar%in to the present day. Cambridge. 5;. 8 Dosenberg J.D. ;??5. The e$olution o# modern human childbirth. Am . *hysical Anthropology :9" p89H1;G. 5>. 8 !riedlander" -anc& K Lordan" Da$id J. 1995. *bstetric implications o# -eanderthal robusticit& and bone densit&. !uman 'volution 5!lorence6 = >>19 >G;. 5G. 8 .iller" Eeo##re& ;??C. <rain e$olution. In Eangestad (.8. and (impson L.0. 5eds6 The evolution o# mind- #undamental .uestions and controversies. Euild#ord. 55. 8 %u)le&" Lulian 19G;. 'volution the modern synthesis. 0llen K =nwin" ,ondon. pG1C 5F. 8 %uskins C.,. 19>1. The origin o# +partina to%nsendii. 1enetica +*" 5>1. 5C. 8 ,amoreu) 8.! and %utt !.<. 19>9. <reed di##erences in resistance to a de#icienc& in $itamin <1 in the #owl. . Agric. /es. 3ashington 9>" >?CH>15. 58. V a b ADob4hansk& T.B 1981. $ob,hans)y2s genetics o# natural populations. eds ,ewontin DC" .oore L0" 3ro$ine 8< and 8allace <. Columbia =ni$ersit& 3ress -.U. 59. 8 Egdar !. 0llin and Lames 0. %opson 199;. E$olution o# the auditor& s&stem in (&napsida 5/.ammal9like reptiles/ and primiti$e mammals6 as seen in the #ossil record. (ection I2 5.ammals6" Chapter ;8" pages 58C9F1G in The evolutionary biology o# hearing edited b& Douglas <. 8ebster" Dichard D. !a&" and 0rthur -. 3opper. (pringer92erlag. I(<- ?9>8C99C58898. F?. 8 -eil (hubin ;??8. :our ;nner Fish- a <ourney into the =.78billion8year history o# the human body 3antheon <ooks ;??8. I(<- 9C89?9>C59G;GGC9;. Chapter 1?" /Ears/ F1. 8 3anchen" 0lec. 199;. Classi#ication, evolution and the nature o# biology. Cambridge. Chapter G %omolog& and the e$idence #or e$olution.

F;. 8 Eould" (tephen La& and Eli4abeth (. 2rba 198;. E)aptation H a missing term in the science o# #orm. *aleobiology >" 1" GH15. F>. 8 8right" (ewall 19>;. The roles o# mutation" inbreeding" crossbreeding" and selection in e$olution. In *roceedings o# the +i"th ;nternational Congress on 1enetics" p>55H>FF. FG. 8 Charles Darwin was the #irst to put #orward such ideas <arrett 3.%. 5ed6 198C. Charles $ar%in2s noteboo)s 518>FH18GG6. Cambridge. F5. 8 2an 2alen ,. 19C>. 0 new e$olutionar& law. 'volutionary Theory +" 1H>?. FF. 8 Darwin in the (rigin o# +pecies tells the stor& o# /a web o# comple) relations/ in$ol$ing heartsease 50iola tricolor6" red clo$er 5Tri#olium pratense" humble9bees 5bumblebees6" mice and cats. (rigin" Fth edition" p5C. FC. 8 Joh" ,ian 3ih. ;??G. +cience" :;9" 5F9?" 1F>;91F>G" 1? (eptember ;??G. F8. 8 3rice TD" Q$arnstrXm 0 K Irwin DE ;??>. The role o# phenot&pic plasticit& in dri$ing genetic e$olution. *roc. Biol. +ci. *?; p1G>>H1GG?. F9. 8 3rice T.D. ;??F. 3henot&pic plasticit&" se)ual selection and the e$olution o# colour patterns. '"p Biol. *;= p;>F8H;>CF C?. 8 .a&nard (mith L. 199>. The theory o# evolution. Cambridge. >rd ed" p>>. C1. 8 .oore ,orna E. and Degensteiner Ludith E. 198>. 0daptation to high altitude. Ann. /ev. Anthropology +*" p;85H>?G. C;. 8 .a&nard (mith uses the term physiologically versatile #or such animals. .a&nard (mith L. 199>. The theory o# evolution. Cambridge. >rd ed" p>;. C>. 8 (ober" Elliott 199>. *hilosophy o# biology. *)#ord. p85H8F CG. 8 8illiams" Eeorge C. 19FF. Adaptation and natural selection- a criti.ue o# some current evolutionary thought. 3rinceton. p8H1? C5. 8 /The h&pothesis that adaptations arise without the e)istence o# a prior purpose" but b& chance ma& change the #itness o# an organism./ ("#ord $ictionary o# >oology. <ut one might :uestion the word chance" since natural selection" b& its operation in particular habitats" is not a random process 5it ma& be a stochastic or probabilistic process" howe$er6. CF. 8 3ittendrigh C.(. 1958. 0daptation" natural selection and beha$ior. In 0. Doe and Eeorge Ea&lord (impson 5eds6 Behavior and evolution. Uale. CC. 8 .a&r" Ernst 19F5. Cause and e##ect in biolog&. In D. ,erner 5ed6 Cause and e##ect. !ree 3ress" -ew Uork. p>>H5?. C8. 8 .a&r" Ernst 1988. To%ard a ne% philosophy o# biology. Chapter > /The multiple meanings o# teleological/. C9. 8 8illiams" Eeorge C. 19FF. Adaptation and natural selection? a criti.ue o# some current evolutionary thought. Chapter 9. 3rinceton. 8?. 8 .onod" Lac:ues 19C1. Chance and necessity- an essay on the natural philosophy o# modern biology. Jnop#" -ew Uork. I(<- ?9>9G9GFF159; 81. 8 -agel" E. 19CC. Teleolog& re$isited goal9directed processes in biolog&. ournal o# *hilosophy ?, ;F1H>?1. 8;. 8 %ull D. ,. 1981. 3hilosoph& and biolog&. In E. !lYistad 5ed6 *hilosophy o# +cience -ijho##.