cpp © 1972 by Ad Pobiting Congas Alstyne No prof tl penton may ‘road ar ented ey foro ‘sero ec el sen ie son neem bye Congres Ctl Number 10405510 Prine inte Utd Ste of Amin w Preface \When Charles Darwin publi his book The Descent of Man and See ton in Reladon to Sex in 1871, be woe in he ncodution: ising ny yar I ctesed noe on te vgn or decent of mas, without ay intention OF pblihing othe abet tu aes withthe deterinton ‘ot to publ sek hohe tat (sboall iur on add he pres ered fo me sflent fo nde, inthe fst elon tiny “vino Sete” tat by thi work “ght woud bs throw en the hig of man and is stor and 8 Ups tht man ens be faced ‘th ofte oreane Denar any general cocasonrespeing hit mone: Ue topearanceon ther. ‘When the late of opinion tome ls profil to the theory of eve tion, Darwin ptheredtogeber his notes and prepared them fo pubes ‘on in Book form. He sates tat his cbservations began in 1898 abd from then on he “ocenionilyatseded fo he subject.” "When prepared fr publica, the nots were voluminous and fll ato cee proupa; those pertaining 19 mas descent, these clang 40 sex ‘elston and thovetelating fo tho expesion ofthe eres Inman en [nial These two tops formed the to past oF hs 1871 book, end His ‘Shnecations onthe eveluton af human exresons llowed i 1873 ‘Darwin's chap onthe decent of man are based na review of ea- tral compacting anatomy and peyhoboy. They do not depend on the tvidence of ay “ining link” i sa’s genealogy. On this bas, which t- fay we would conser very line, Dsrwa reached surprisingly ssurate Gotsasions es fo ats sacetor (en early Catarhing led to the gorila Sd cimparae), be Bplay (a bot county, probably Aice), an the Time of hit origin (between the Becene and Miocene). Hi views were ‘love to thowo of EretFoeskl,pubabed la 1868 undsc the thle Narr lite Schéplonggeshichi, aithoueh they Were developed independently. oman evolution had ben eicesed by Haley (Mans Plce In Nature) i7 Parental Investment and Sexual Selection Iriroduction (Chale Doewi's (1871) Ueatmant ofthe tople of sexual selection was Sometinos confused becauue be lacked a georal framework wii which ‘ores the varibls he peresved to be importa: sxlakedlnbecanes, ‘ax ratio at concepdon,diferental mori, parental ire, and the xm the breading ater (monognny, plyayny, polyaniy, or promiscuity). “This contusion permite others oatempe to show that Dar’ temil- ony was impress, that he mbaterpeted the futon of some sueares, td that the Influence of sexe) selection wat reatiy vera Holey (1938), for exampo, dsalser the impoctsnce of fetnale hole without Dridenc or theoretic! argument, and he doubl the prevalence of adap ons in male tat decease thst chances ct rriving Sut ae slected be. aise they lead to high reproductive suszess. Some importat advances, Foweve, have been sebeved vege Darn’ work."Te geet of sex now been clare, and Fisher (1958) his produced a mods to expan ex ce at concepion a model rcenly eanded to ncude special mecha ‘ss that operate unde inbreeding (Harton 1967), Data fon: he lb ratory and the Be have confirmed hat emo ae capable of very suble Soles (for example, Pe Ehrman 1969), and Batewsan (1948) bas sug ‘Suted gunerl brat for female cole aed malesmale competion, snd oa produced peels data os oe spel fo suppor i argent ' Sehoeer for sos ction Thank Qf. Suderned (Harvra Seats apr ie a oy me ae we EE Milam een dient ad wna seat eso The we Parental Invesinent and Sen Selection 137 ‘hie paper presents « general framework within whleh to consider sexual scion. Iai Tatempe io dine und Inerelae the key variables. [No atcmpt i made to review the lage, steed Iecatretelevat to ferual selection, Instead, arguments are presened oa how one might expec aataa!elestion to aot onthe sete, apd tome data are presented ‘oauppor thse argument, Verlance tn Reproductive Success Darwin defined sex! selection as (1) competion wih one se for member af the opponte sex an (2) diferent! choice by merbers of 008 Se for members ofthe oppose sx, and he pointed out thst this usally meant males comping wits each efor for fmales and females choosing soma males rater that others. To xudy Uns phenomena one needs ao fcurste dats on diterenn reproductive sucess agalyaed by sex. Accurato data‘ fence reproduce socom ae svalebe for many opocies, Dut ‘lr data on mals ar very cl to gue, even fa those spect that tend toward menogan, The human spies ltrs this pol. In any society itis reaivaly easy to esign actly the hire tothe blog ical mothers, but an element of uncertainty ataches tothe usignment ot children to thle bloga! father. For example, Heny Harpending (p= ena! comemunicstion) fas gathered blochemial data on the ‘Kalba Bushmen showing tat about two por cent athe ehldrea In tha sekety do not belong tothe father to whom they ae commonly atlbutd. Dat onthe homie species ae, of cour, mach more dtalled than sae d lon other species. “To gather precise dats on bth sexes Bateman (1948) suid a singe species, Drosophila melanogaster, undet laboratory conditions. By sing chromosomally marked inven in competion vith individosl beat ing dierent marers, and by searing forthe markers in the ofspeng he vs able to measure the reproduce ace of each fli, whether female or ral. His method conseed of intedcing five st ales to five adult fea veg, so tht each female hed a choise of fe males and cach male competed with for ote males, ‘Dts fom numerous campeiion experiments with Drosophila revealed thee important sexual difereces: (1) Male rprodustive succes varied uch more widely tha fomale reproductive success, Oly four per cent ofthe females led to produce any surviving offspring, wile 21 per cent ‘ofthe mnles so fas, Some mas, on the otker Bend, ere pbenemesaly ‘sce, producing nearly hres tines ax many offspring atthe moe e+ ‘sual fru. (2) Female reprogotve sucess did nt appear to be in ied by ability to atract male. The four per ceat who fed to copsate were apparently courted as vigorouly ak those who did copulate. Oa the bts Rand, male repoduatve sucess was severely Heed by ably toa ones suvens strat or aroun forues, The 21 per cnt who fled to reproduce shoved to direst in trying to copulate only un lblty to bo aczeptee. (3) A Females seprodustive acon didnot increase ech, i any, after the ft copulation and no at all alr the second; mos females were uninterested In opalatiag more chun ence or tele, As show by gente markers nthe sflping rls showed an aostUeatintese la reproductive sinest ith Iaeeasedcopulations, (A corlazy of the fading Je tat malar Ietded sotto mate withthe soos ferla twice.) Although these rele Weve bisze in the laboratory, they ray apply with cven greater force {othe wid, where mae are med to fe females und Woe females Inve wer rango of males fom whlch oehooe ‘Beeman argued tat hi ela could be expuined by reference tothe enogyinvesteat of eth sx intel ex eel, Slceeale Drosophila ine test very litle metal eeery in the prodction of x given tex ely whereas females Ines considerable energy, a male's reproduce sucess Is not Haste by hls ity to produce nex cls But by bilty 10 fete [as egg with thse eas A frale's reproductive wuscese i tlie by er ay to ae Ber opp fetlzd bat by her ably to produce eae ‘Since in elmo all eloal and pleat spelt the male produces sox cals that tny by compution to the font’ sex cells Beran (1948) sued that his results should apely very widely, tht is to “all bate few Very primitive organs, and those la which monogamy combined th ‘ext of wily elininstd al etree lection” (Good Sil data‘ roproducie seen re iit to fad, bat what dain ein Snjaeetion withthe sotopton that ale reproductive na fess vara 8 fonction of he mamber of copultions,* support the conten. tion that inal pete, excep hose metoned below in which male pan ta cae may be aiming espace for fama, mae repedvtive see ‘aries mere than femal reproductive sucess. "Tis is supported, forex tmple, by data from dragonies scabs 1955), baboone (DeVere 1965), omnmon fogs (Sevage 1961), parle chickens (Robel 1966), sage grow (Soot 1942), lack rouse (Koivisto 1965), elephant seals (LeBoeut & Petron, 1969), cur ies (Pater 1970s) and sme anon ards (Rand 1967 and Tives, in preparation, disused below.) Cheunstinialev- dance exits for oir ards (or exams Bair 1960, Haris 1964) and for many pammals (se Eiterberg 1965). Im monopnnous speci, male reproduce sucess would be expected to vary as female reproductive foes, bit there i lays the pomibily of adahery and ferential fe rule morally (aleused below) and these factors shold Ineease che tat ie fone srl ng Sth ee ana, abo tae {eben eampl atonan Sah Senge 66h Boras 968 bt nto Prk ED). Parental Invetment and Seal Selecson 19 Yariase of mole rpreducive succes without sigaicntly altering that oth fone, Relaive Pernt tnvetment Baten’ argument can be sated in a more peels and genta form sch that the breeding sytem (for erample,monogariy) at wal a the Adult sx ratio Become functions ofa single variable cotroling seal lection. I fst define parental inerimentat any Inverinen BY the parent In am individual oftpring tht inves the ofeprng's chance of sting (and hence reproductive sueces) othe cost ofthe parent al fae Yt n other ofsping. So dene, parent investment inltethe tet boli investment inthe pray sex ale but rere fo any lesen (sch 1 fending or guarding the young) that beoeis the young. Te oes nt fe clude effort expeaded in fading * member ofthe eppoite sex ori sb ing members of one's own tx in odes 9 mae wth a member cf the Op peste ser, since suck efor (excep in spcal cases) does pot afer uve chances of te esalig ofspring and is Uevefore not parent investnet, Each ofpring canbe vewes stan ievestment independent of ater of spring, Increasing invenent Ia one ofpring tending to doucate ine: tent in other, Irmeasute the sze of = paventl invest by selene to is nepative eect on the parents aby To fnvext in oerofpelng arp parental investment Isobe that strongly decrees the paren sbity fo Prodoce other afuping. Thre eno neneary covrlon between the se ft parental aves nan offspring and ls benett forthe young Teed, ‘ns cin show tht dering s breeding season the Snell fmm a Bvt [ateatllvestnent mst deren some point or ele specs woul at ‘endo produce any Bned number of spring ger ston, Desa in fe producve muses esl from the cogtve lle! of prea iver ‘ent on nonprental forms of eprodstve effort (such 8s sera compet tin fr mata) Is exloded fom the tesrorement of parcotlinvestoes, In eles, then, I ar bere consering productive uozas ae Ht On) relevant lable wer pasa aves Fora giten reproductive sean ope cn die the ttl parental invest ret ofan odvidual athe sum of fvestnents In each ot png rodcsed curing that sason, an one sues that tral sections hes favored the total parete nvestnem thet lade Yo mex ne reper doctive succes. Dividing the total parental investment by the umber af individuals produeed by the parent gives the Ipical parental inesteest by lll! per orig. Batemsa's argument ean now be rear Ite as folios, Sins the tte mmr of asi produced by one se fa sexually reproducing specs mest equal the toa number proded by the othe (and sasuming the snes Ase ino other Way than In thle0 omexe evans RS or PI ™ L NO. of OFFSPRING PRODUCED Pre 71, Reroasn uct (RS) ad dct in tie rpc ‘zs renthing fom pare Ietimet CE) re raphe oe lerctot of the Mune of aBpring prac bya of {iia Mah regi he rer ‘nasi fo er bond ee 9 rpetvy. Se? ond 8 Ge tah. The sope of the cuves need wo be specfed xs. ‘pial petal invent per ofa? then the sex whow eat puro ivetnent pet! han ato he oppose sex wil became Aehtng essre fc ht elds fh avenge wl om ‘hong this wo rod ith ener € the et hvetng 3, {ean tna of he forme an lneee prods eo By inesing ueceey inte oping of veal member of Tinie {20 By mag & ape fcoahp beeen dope of ctl inv that and tumir ol oloping product, angst canbe preted teabily (igure 71)" pti for sexe competion he tx Aicatng les etn be ead by ealeltig the ofthe ember of pig hat ex optimally rokcer (a fusion paral inves rey 5/5 x cen an (baer andre mony yr ea ator ca raat sek rao, as ao be rete wfc of seal ‘Boston Fe spect pry a pei een es ee $3 et har he uigtie ee cn eet Puree! Ievenment od Ser Seles ua ment alos, assuming te oppose sexs ivestneat fed atts opine vale) to the aumber of offspring the liming ex optialy produces Min Figue 7.) ‘What governs the operation of serualslecton & the relative prenal lnvestnen ofthe sexes Tn ter ofsring, Competiion fo mates taal characters males becaue mates mally Havest almost nothing in thir ‘pring. Where mule parental invetnent per ofping ie cxiparsble to fe rnle investment one would expect male and female repodacive suce to vary sinfar mays and for feral cole to be no mate dleeinisatng than tale choise” Ceacept at noted below). Where stale parental i vesnent stony exces that ofthe femnle (egadns of which ae le ‘ests more in the sex eal) one wold expect females fo compete song hares for mals and for males tobe slestive abou whom they accept ‘Nat tht it may not be posible fora iil of one sx o invest in cay pat ofthe offspring ofan individual of the epposte sex. When Invests less pe typealllspring tan dos a female bat moe Gan on hall what she invents (or visors) theo section may 0% faer mie compe ‘ton to pit with move then os fle, I he ofipring of te wosond fe ‘ale canot be pucled out to more thas one malt the net reprodae™ ve seces for a mae investing Inthe ofrag of one foal else han that guined fom Investing in the oping of Ivo females, hes the sale wil be selected to inves in the ofsplag of ony one female, Tas ‘segment i graphed in Figure 7.2 end may be isprtne vo aodertanding Aifetemialrtliy in monogamous bed, 2s dcused elo. (1958) sax rato model compare the parent expenltare (un- ened) ia mle ofting with that i onleaaping and svegests energy fend tine as measures of expendlur, Resatoments of Fibers model (lor ‘ample, Kelman 1960, Wileos & Blanks 1963, Emlen 1968, Verses 1963, Leigh 1970) erly ete he undeined term, pzetal expendi, rte term energy investen. I lhe cage the ey cone impels tnd the relevant one is pareatllvesinen, at defined above, Boer i ‘wesimené may often be & good eppeoximation of parexalivestnt, bit ‘tis clerysemetiner& poor one. An indus! defending ite bod fom ' predator may expend very ile eneray In the process ttle high hance of mortaliy such beavie should be menaced as 2 Ite Inver ‘ant not sal on wget bythe ener ined Pavel InveximentPaternt ‘Species can be easied sesrdag to the relative parental lveximent of te sexes in hee young In tha vast majority of apes, the mle oly ontribation othe survival of his ofsring i be aex cel hee species, fara contribution ay exeenda ele ae by rg ati, 'A male may invest ins ofprng in several ways. He may provide hsie sooeer & rvans RS (EE conorion 0) nese RS or PI (SE%'2,conorrion ay (EEE conomon or N 2N NO. of OFFSPRING PRODUCED Pure 72. RS and Pa uncon oth mba of obtrngprodyced jor oe Sxay. Set 2 vena gor tpl ofepring more thon hal 97 het iment Condon A! mann nt RS ora enero} Pezming ea st tn any tr of baying beteeen Gn 2N. Condon Bnet RS arming mer of oe? bet aN oping Contin C2 et RS semng member of et 3 van iW obit If member of see ma ive ‘cpa multe of ¥ orig, natal ein vrs conan C. ‘mato with fod asin aloon fles (Kesel 1935) and some other insees (Geguimann 1970), some spides, and some bids (ler example, Cale 1967, Royirs 1966, Stokoe & Wilms, 1971). He may nd std defend po ple forthe female to fed ny ogo ase young as ln mn bids fie may bul nest to receive the pg, ei Soma Bah (lor exampl Moai 1952), He may help the female ly te ets a a some pasate birds (Lack 1968), The male may elso defend the fesmle, He may Dood the eng, a in some bits, sh, fogs, sed slamenéers. He may help feed the young, pretee them, provide opportune fr learalg, at 0 Go, a8 im wolves and many monogamous tres. Fall, he may provide sie let group beset to the young (such a protien), afin may pi mates. Al ofthese forms of mle parental investment tnd to decreate the tard inva from sch tk. Tt eu ose hows mal cule be vocal in deceiving n ne fol, but i a female ats oll, she right foot a male. Ar tine goes on {for example, nce the eg ce Id),‘tually thet a ale cul easily Be fle. The female could thus be Programmed to try the hid etategy fine and tae, to rvet tthe First or seconds Theta dex gins oti the female soccoods fn the third statgy, noting it she chose the Ba sag), and possibly an Intermediate vale I sho chooses the second rae, eer parece desert at he Begining, then a te goes on, exch tn vests mors si ore the young Ti rend his several consequeacss, the one hand the pir of deter more capable of isl the task ‘lone no natral elation shoal avr Hs being more predisposed (0 (9, becouse it hes more f0 fre, On the ether hat the dessa more to Tove if he poring flan least glo the pavtace succeeds, The bale face between these opposig Ettore sboul depend on te exact form of the comunte investment cues at ell asthe opportunites fr further breeding outs the pale “Trove another ees with tine of the Lacesing fvestset by both parent Inthe offspring. Ar he fventnets ise, antral selection say Favor either prterdeseing een i ope has nested more inthe young than te other, hi ls bea the deveton sey put the deserted pera int crt binds he bs invested ro sh that be loses conserby i he tlio detects the young, en though which should make no aiference to ‘hin, the parte would lowe een more The polity of seh binds em be fused by an analogous station desetbed by Rowley (1965). Two ‘eihboring pro waeeshappesed to fede tne young slltanectly od could ot tel ther young aper20 Both pe eal sb Youn ics- ‘tmiantely, unl ene pale "deserted to ris anther brood, leaving tsi Seber fo fod ll sx young, whih they €, even thogh thls meant ‘hy wee, eflec, bagten advantage o- ‘ire should show edepsions to avod being deere. Females, a pac tial shoul be let guard agnet males who wil oxy copie nd Dot invest aubeogent parental eff. AB insane of such an adaptation tiny be found inthe re-necked plate, Ploropar lobar. In pale ‘opos the male nce te epg alone ad alone ears for the young afer hashing (Hin 1867, Johns 1969), so hat s prop of eumuatve parental feveatent would show an fia Tage femate invests which then = ‘in the sme though tie, wheres the aia male invesimest iol od incase stl, probably to surpaethe fale lesen. Only the female ylnrebe to being deserted and this ght after copulation, shee ‘any ler dveton bythe male cons him Bs lvestnet In toubato, the {young being sans ortsin to perish Tinbergen (1935) chert female Yinorvsy courting ale and then Sag avay as soca as he reponded Tove courtship by attempting to coplte, Ts eoy performance Was = pated mamerots ter for sveral ys. Tinbergen attbuted i to the Frog and waning ofa atin” but the behavior soy have been et ereilInvetinet an See! Seen v9 cof the male's willgness fo brood the fomalé's egg. The male under ober Srsion was, in fac seedy roodlog ogg thd was courted whee be ltt theese to feed om a nenby pon, Ta rte fo view a complet egedaing feence, Tinbergen destroyed te euch the male was brooding Win TR dy the fecsle permed te male seal acess, and he subse {uenlybrooded her eggs The Important pot Is that the fale could ppucnty we the deren bewenn a fre and an encumbered mal, and {Re wtheld sex fom the later Coutsip alternating with Might may be ‘he test tha eval the mul’ tre tachment the ts ean Hb, oF eX tmp heer eo fallow the fama. Tris Ikaly that many aGepations exis ls monognous sete goad sgsnst desedion but despite evidence thet dereion can be common (lowly 1965) noone bas stems lo analyze courtship with his danger Si mind, Von Hantiman (1965) tng reviewed some cvleny for adpta- tion of feats to avoid Song mated to a polygynos rma, ndbeing 20 ‘aad is sometine easly eqlvalnt ro beng seria by thy male (von Haaren, 1951). ‘Eterm feiation rogues a synchrony of bebavin sh tat the male can wily be certain he i Rot atempting to fer, previously {enon eggs With the evolston of itera etizton the ale camnct tbvo cet For many sens (or example, mest mama), the dstne- tin set importatbectse the mal oes oil by stenting to fet fee previously feriiaed gar. Where male parcial cei ive, bow (rere mate cons the i of being evcklded of raking another ale’s ‘Sipdng, Por Figre 7. twat astred that the pale copaatd with each ‘ther and each eter ony, but the male can usually not-be sue that ch frie cae and what roped in ch station [s the males ipestment Ink femal’ ofeping, Adapttons Should evelve to help guarantee that the female's oping ave als he ohm, but cose con pry be couteed bythe eveluton of more sophisicatedcockols ‘One way a mele can pect inself fo ensure that oer males Seep tna Stace. That some estos) agpresion of monogamocsmale bids Is devoted to protesting the snc ofthe pair bond ems cera, ard e- fran mnie agyresion toward real Or sapeted dultees Is often etree {ee (1969) for example, ba shown tha, when the case Is Known, the ‘arene of tel Buthoan fh x adultery or uapectd dues. Tn ited data on other buncrsbecng groope (ncodig Esksice and ‘Attain aborts) indlone that, while fighting is celatvely are (in that orgasaed intergroup aggstion ie atequent), the “mrder ete” may te vltel high, Oa examination the murderer and hs visi ae usualy fhusbnd and is wiles reat or suspected lover. tn pigeons (Columba livia) anew mae acving lone a woctural roosting pace in the fall it ‘Machel day afer day by ons or mor aséeot ras, AB 4000 a te sameraale appears with a mate, the two ae trated much more cosy (rivers, unpublted data), sugering that an unpived male more ‘Sretening than pared one Tave argued ove that fmgle deserted lnumeclately eter copol tion may be adgted to ty Yo fdce another male to belp rae hee young "This tacor imple ndapatione onthe part of he mle to eld such se ‘A dngle meth leo weld eating with & feral on Set ensounter, 98 ‘que er lstend and mate with er ony afer a pssege of fie that res {onthly excludes oe por Inpregnation by soother mal, Cea males ‘hard tholfeale fam ote mals, end thece 9 stcking ference be- tee th lack of pinion In promizeuws bie (Sot 1942, Keit & Hogan 1967) andthe sometimes lng ag between pic bonding and cops luton ia manogarocs bird (Nevo 1956), «lag whi uml seems to tere fer funtion wel ‘Bilogt have Interpreted courtship ins mlted way. Courtihip is seo ss allowing the individual to chose the correct species an tex 10 ove ome anagoitc ares and to aoe one patter (Bastozk 1967). The shove aalyee sugges that courship should alo be infeed in ira ‘af the need to gard oneet rom he several pss of malieaest at ‘Bebande of one mate Dire! Mortliy andthe Sex Ratio f special interest in undersanding the efi of seul setion are 3 corte data on aifeeadal morality of the sbes,eepdialiy of fmature Tnaividos, Such data ar, however, anong the most dat to gather tnd the publshed da, although important, are seanty (for example Elon 1940, Heys 1947, Chapan, Cosi, & Cote 1938, Robina ee. 1957, Coulan 1960, Pts 1969, Diag 1971, Myers & Krebs 1971). A tttute one can a tte of dats on vx ratios wihin given age cle fr forall age cates ken logeter. By sssuning thatthe ex rao cote ception (oe, les pct, at birth) ts almost exatiy $0/50, sigaiant evans thom this rato for any ae class or forall taken togtber should Spl diferenl tray. Where dats ext for the sex cao at bith aod her the sox ratio for the entkeloel population ir unbalanced, the sex aoa ithe unully about 50/50 (se above retreees, Seandee 1965, Tack 1954). Parthemore, Fiber (1958) har shown, and others eased (Leigh 1970), that paren shold Saves oughly exual energy i eah sex ‘Sie parete'sually invest roughly equal energy in ech iaividun of ‘ach st, matt seleton In the absence of unurul cumstances (230 anioe 1967), sold favor apres «30/50 x rao cote ception. Parental Invenant and See Secon 1s ts ifet to dtrine szuratey the sex ratio for any species, Te ros serous souece of bine i tnt oes and feoales of make the {ies difercately sya tothe err or example, in soll msm tls seal selection seme to have favored mle abt, soc a igh ‘nob, that tend fo esl hele deren capture (Bee, Prone & MacLecd 1958; Myerr & Krebs, 1971), Tone views on’ eaptuetech- niques as iaadomiy stmpling the exstag population, coe wil eoodude {Gat males ae move momeroe, IC one views ooo capture febriques at randomly sampling the efeets of mortally on the poplaon, then ons Wil conta that males are more pre fo morality (Ihe are expared Ine often) aed therefore oe ler murecous. Neither assumption is Hal tobe tr, but authors rtnely choose the former. Furthetmar, ke fea Sot appreciated wint «lage sompe rogied fn onder to show dine ine devitons tom = 50/50 rato. A sample of 400 aninals showing & 41/56 we ratio, for example, doesnot deviate sgalcaniy from a $0/50 fio. (Now ahough ths ios neverpolted ext, does i lifer sige tent fom 238/62 ral.) ‘Mayr (1983) fas polated out tht there are nuserous deviations from 450/60 sen rat ke bide and X lve Wt Ue that if dat were Saflenty prose, sat apes of vertcbtze would show a sgalfeant evan for » 30/30 sex eto, Nan and females dir In numerous ‘Surectrisicy relevant to thelr diferent reproductive sities nd thee ‘hasctr ar snely to ave eqamtet fects survive, Slo tis et advantageous forthe adals ot eae sex to have avalabe the same umber of ads ofthe opposite se, hee wil be no astra skecive tent for kepng cevitone om a $0/50 ratio sal A veview ofthe wel Hertre on Sx tabs suggaats that (except for beds) when the sector onbleneed Ke wualy unbalanced by Phare bring more fess than male, Pat another way, mals appareatiy have {tendeney fo sifer higher morety rates than feoles, This i tre for those dragons for wbih thre ae data (Covéet, Longfield, te Moore 1960), forte note By (Rockatsn 1959), for mest Bs (Beveton & elt 1939) fr erea inde (Tikle 1967, Hares 1964, Hirth 1962, Blair 1960, ‘Teves, dicted below) and for many mammals (Boule & Ver sehuren 1960, Cowan 1950, Ezenberg 1965, Robnetie etal, 1957, Bes Fenecl, & Macleod 1958, tphene 1952, Tyndale Bisoe & Smith 1983, Myers & Krein, 1971, Wood 1970), Sailln (1948) ad Lack (1958) have revewed saves on other animals soggesing a simi tend. Maye G30) points out that where the sex rato can be shown tobe unblaned tn monogamons birds thre ae stl fewer fetes, bat npoygyaous oF frolocuus bids tte ae fewer males. Data since his paper con this Fnaing Tats rest is patclry Interesting slsce 1 all other groups fo ‘rich mals tad fo bees trou monogany i ee or onexisent.is owen riven There a tendency among Viegas study sola behavior to regard ‘he alt sx rato asa independent viable to which the specs sets ‘th eppropriate adaptations. Lack (1968) ote interprets roi behavior tr an alopttion in pat to an unbalanced (or balsosd) sex zat, apd ‘Verner (1964) has semmarind other instacen of hs tedency. The oly Inechinism that wil generte ferential morkliy indepeadeat of sexual Aiferences cles rete to parental invetnent and sexual sletion Is the choriosoml mechanim, applied expetly to humans and other ‘mana the unguarded camorene ofthe male is pesuned to pre- ‘depore hi to higher moral. The mechani le inadeguate asa ex- nation of eierentit mortality for tres reson. The aaibtin of eifarentisl mortally by sox ie not predicted by © vowadge of the dbution of sex determining, mechanisms. Both sexes of fob ar usualy homogamet, yt mass sue higher moraliy. Female binds are beterogametic bat ser higher mort ony in monognmoas specks. Homognmetie male meal athe are ostvived hy tier hetero. fmt female coanterpars vader laberatry condone (Haniton & Seareson 1965) 2. Toeoretial preditions of the depco of difereaal moray ox led by males dbs to tel unguarded X clemson are fr Lower than Those observed in soch mamas a dogs, cate and hus (Lodwlg & Boost 1981). Tels pose iagine aural selection faving the beter ‘ow sex determining mechan i the aseined srt) mortality [light aod balanced by some advantage in ifeentiation ori he homo fume sox, but large morality ancised with heterogemy show be counteracted by a tendency toward both sexes becoming Romogametic. 3, Corfe ata for humans demosszue that caste males (Who remain of coors beterogamete) stony outurvive conta group of Tole sila inal eer respess and te aie in ie the easton, the eater the increase in surviea. (Hamilton & Mesie 1969). The sare is tru of domestc cats (Hamfton, Hamiton & Mester 1969), bot not of @ species (meal moth) for which tere fr no evidence that te gone are Soplated in sexual ferentiation (Hamion& Fhansso 1963) ‘An Adaptive Model of Digerentiel Morsay ‘Tointerpret the meaning o talanced or unbalanced eet ros one needs a compretensiveframewerk within whith to view lie historic! phenom: fa, Greil & Bosct (1970) have presented a model for the adapve Intezpreation of difeencs beweenspeie lie stories; for example, in the ag af fst breding and in the growth and survival curves. Although they eid not apply te model to real llerences in thee purest, Parent Invenient and Seal Seton is the modell preiely sited for mch dBerences. Ose ca, in eflet, teat the sees af they were cferet speci, te oppose ex being 2 retouee sant to produng ximsm surviving ofieag. Put hs way, fea pies” uly ier fom mle epeir in tat females compete among themes for och reioures as fod but net for mane ofthe oppose fou wheres fs alstely compste only for members ef th oppose fx all ter forse of competion Belg important oly isoar es they af fee thi alinat empaien, "To analet dierent mortally by sex one needs wo corlate direst reproductive sratgin with moval, tht fy one must sow How @ gen ‘predacive astegy ental given se of morality. One can do this by irphing reproductive sucess (RS) fr the rst reeingsson es «fone lon of reprodetive efor expended during that seston and by graphing the inination i ute reproductive even (D) fn unite of st breeding season reprogatve asses. (Gadgl and Bosert show tha te epredue- tive moe of a given ello doles ith age, hence tho ned to comer. fate reproductive sucess to comparsle wai.) For sxpley assume thatthe diminution, D, resus esti rom morality between the At and Second breding seme The dnlatlon coud emi from moa fa 8 Inter yer (indooed by reprodotive effort ia the fist breeding esta) hich would not change the form ole analy, o could rr eon de- ‘sensed billy t0 brad inthe second (or tll ler) breeding vesson, ‘which sometimes cece But which & probably minor compar to the miaton dus to morality, aod whch dees at change the anaes feng one stsumes tat male and females do not Aller appreciably inthe ‘xt which they afer th for of insti. ‘Nara selon favor an indian! expending la cho Sst breeding season the reproductive efor (RE) that rel a 8 maniurn net epcor Get seems (RED), The vs of D atthe RE gies he dapee of ex Deseo motaliy between the ft and second. breedlg essons (see Figures 7a and 73), Diereoesbstneen the wnet in D wl give the ex: peste éideentia! metal. The same analyse can be appled to te a Freeing season fo predict snort between i snd the ath + 1 breeding cvon, Likes, bys tal modieaton, the als can be sed To gear trate dferenes In avenle mori: let D reproveat the diminution in hanes of surviving fo the Sat breeding semen ata faction of RE at frst fone ie easing the coin survival of dewelop- {oad reproductive In Figure 7, have graphed RS and D as fnetlons of sepraductive elo in the fist breeding season fc fener of 8 hypotblcal species Is whichrile intest very ite patel care. The RS fonton i givens sgmeldsl thape forthe folowing reasons. Tasmume tat at low vales of RE, RS in reas ool) ery gradually Beonee some investmeat is eseeiy ut inne reproduction (or example enlarging the reproduce orgs). RS then ineente more reply st» Rmoton of RE bot withon ahiving # vey Heap slope, RS aly lvls oft igh values of RE Beetee of Ine (retved foeficlecies there (for example, inefelenses in fag Schoener 1971). Thavegrapbed the vl, at whieh ne reproductive soe- ‘est fr the female veaces « maximum, Tecnlaly, de fo ompetton, the shape of the RS fanetion fr any given female wil depend pany on tie ropeeductis effort devoted by eter females; the graph therefore as tes that oer females tend to ives near the opal wef, but Important festre ofa female's RS i tht I snot ronly dependent on the RE devoted by oer females the ourve would not geal ier Ia bee female invested mach more or les. T have grapbed D at a neat funetion of RE. So doing anaunts to a defniton of reproduce ofr, That 0 given Icrement in vepoductve effort during the Bat breeding season ean be deteced as a propronatly increased chance of dyleg ber female Net REPRODUCTIVE SUCCESS, RS and 0 f REPRODUCTIVE EFFORT Fiqwre 74, Female eposucve maces daring the ft beeing recon (85) ‘a dian of fur reproducing sates (BY on funsons freqrodutie srt ring fet breeding. mesured Ins 2 fe reg i) eet ae each RS. Peenal Investment and Sexual é | male NET ReeRopuctive success D RS ond D m REPRODUCTIVE EFFORT Pigane 75, Same Bgure 74 xen ha drawn orth mle ied ofthe Somat Atte net reproductive ces racer ¢ masini tween the fist and scond brsting sestons, Note that reproductive efor. forte female is set syaonymon with prea aves "Moe RS efor from fale RS in two important ways, both of which seem rom sexosl election (1) A male’ RS i highly dependent on the RE. tater males, When other male favest best, am indivi mal wll, Chaly ot ovteompts them nls be vet 85 much or more. A com fiderable Investment that ail below that of tbe males may rex [acto RS. (2) A mole's RS potty very Big, uch bighec tha hat ft conspcte female, bot only Ihe eatompetes oter mates. The oud exit some fstor oe wtof factors (rach aan agresivenes, o- ‘iy that covelates thigh male RS. The elles ef eorpatiion te- {wea male for females sleton for ncested mal RE, and tis elec ‘Son wll cominge un reat ale than female RE selected song ashe higher asocoted D isoflet by the ptenillyyery Bit RS. This aq Teens graphed in Figure 7-5, where he seep slope of BS reacts he ih iteration between one ties RS and the RE of te ober males, NO thatthe egumont hee depends te extence ofa tof factors corre ned with high mae rprodutive succes. Tf thew factors ex, oat ‘Selection wl respon the male ohigher morality cates han the feral ‘Whore «mal an ehieve very high HS inn breeding season (a in Tande breeding vet, Bartholomew 1970), diferent mostaliy wl be cor responding i186 sonenr evans The analysis here applies 10 species in which males invest less parental are then Bt probably more then one-half, what fermass ives Tass ‘at most monogamous Bis ae 20 chamcered, and T hive Ised ea sons and some dats above suppntng this asumption, The rezone can be ummrlad by aajag tat Because of thet inal lags investment, fe- ‘ales appese to be caught ina altustion ln whlch they ae nsble to free restr petal lnvertment oot ofthe melee and would be strony {Seed against hey unlateraly reduced thelr own paca ives ‘Funedoas slang RS to pret investment ee grape for males and ‘omales in Figures 7.6 snd 77, asulng fo each sex tht th oppasie sex shows He pareota avesta! tat resus fo it in a manimacn ‘ot Tepo- Auetvesucss, The fame curve i given 8 sigmoidal shape for the ea- soas that apply to Figuce 74 ta Beds the females ina avesment In the gs oil go for aathing if moee b not Ivete in brooding the eggs sd fedlog the young, while beyond a atin high RE furterIncenents do no ray sfc RS. Asumiag the eale faves the value, fale RS ‘vllvay ae fon of mae parental lesen away sie to f= mse RS, exept the funtion wil be dgplacod to het (Figure 7.7) and tome RS wil be let due to the elects of he cackolty graphed in Figo 78 "Beesoe males lav io pretal cate more than eoerhlf what fgsles female logge — Rs RS ond D Re Figure 76, Famaerprodacive scents ne detonate veprobucve ‘casa futon of rpraucve fort (RE) suming ile ‘graduate sont of. Specs hypothe! manage SEC inatck mole et romewhat ee han ema tn parents! fare (ue Fire 7. ond 73, Perel Invent and Sel Sesion 10 male | seer ° ae RS ond 0 my RE (PARENTAL INVESTMENT) Plgre 7.2. Mele rprucin ces and dination t fate sprouse “AG funn of rerouting faa repo theives of Spee ome arn Pre 2. Reprodatoe fer alm td ep car ioe fl cr Invest and because the oping of ven female tend to be inseminated by a single mal, secon doe oot fave tes competing wih each eter to invest inthe ofpring of rece than one female, Retr, sexu seleton ‘nly opertes on the wale fo Inseminat tomales whose oping be wll ot rae, especialy i another male lass them inset Shoe sletion prensmably does no sronly favor female adultery ane may oppo ample, detection lens fo desertion by the mate, the opporsiies for eacksdry ae Unto high ivestect In. promiscuous activity wil bring only lined RS, This argument grphod in Figure 7.8, The pre cid deren moraity by eek ean be ha by somparing D (0) with Di ms) ty tem ironic but in moving trom spromsaovs to & monogsmous Me that, in moving toward greater parent invest In Be young, te tale tends to Unerease his chances of surviving eats tothe feral This tendency ovarebeease tha lncreaseé parental invert droped ely dereases the male's RE lesen malevonle competent ise= tite tae, "Nothin bth eases sbove dees] mortality tends to be mie ing. By altesing the ratio of pose sexual psec to smu competitors Arena! morality sets up foress that tend to keep the dietetistmore tly low. In species showing litle mele penal investmet diereta sale moray iperedses the aeragenasiberof females eveable forthe les who survive. Otber ings being equ he Jacess tend fo make itmore ict forthe most cent males to matin thr eae| male RS ond 0 7 7 ? fe (PROMISCUOUS INVESTMENT) lowe 78. Male reproductive cer ad dination of fue reproductive were te fon of ror artsy devoed fo 970 ‘caus bean Mt reprosuctive ees a a mato Same seta n Puts 3.6 and 7.7. antag. In monepnous bis diferent fale moc ids come pein aon i ef eat om ty ytd ‘Se ole meal. Sch ompetitn Prumaby a Ines te Cave Ia ale epogeste vroae above the sea reat ex poet from ck. the above aunens were mae wh tenet ate pret ites on hey apt pss owiehe ie more Pi fe an oa a wee rae ‘ih muh ean vere moe hn Ol at mle inves ‘one would predict difecental female mortally. Where feriales Invest less tt ow‘ wat mals ave one woud pred compeion, sd re tslegaiferentl fel moray Male-ée Competition neon beeen miles doe no nec cd with he ine of sun "Ere tv apes wih lea feria, compton Between Sfem of tiferet aes at be am inpocast campo of wale-male ‘Breton (teenie by Pater 19708) Tear cae com iton button mls my coms ster egy re tried For x= Parental biveeant and Sera Seeton 159 ample an adult male langur (Presb entlus) wo outs the adult mle of & prop may sstemaiay ll the fans ol tht group (presumably fatbered by Ge cared mile) heey bringing mot of the ele feng thy in estas agin (Sugiyama 1967). Walle clesty dedvantageou orth killed infants end thet other, each bebavin, besetting the male, may be an exteme protuct of sal selection, Few mice spe tanoomly abort during the fist fous daye af pagmancy waen expose fo the sel of trange male (Bruce 1960, reviewed in Sale 1967). ion subject to several interpretations lacing ove based on alo male compan, Spem competition may have importent eects on compton tween smiles peir to rele of sperm. In thaw ints fe which Iteverng ‘em take procedence In Setting eggs, selection fovore mating wih = female just por to relete of eqs, there inretaag competion 2 ort lation sts and Intersting election for postovulstory gud Sto bythe mae (ee Pare 19708, Joube 1953). Ther coneetat ob ule. ‘male compton prior co the release of epi in specie showing Yep lite mate acetate "The form of maleale competion shouldbe strongly ifesced by the slstibaton Jo spice and tine of the ulimatetesureesfeing male fe Productive sucees, namely, conspeito breeding fem, The datcbation ‘be deseled in tems of three parameter: th eet fo whch fete, fe clumped or dispersed in space the extent fo which they ate lumped or dispersed in Ging, ad the ett to which thie enact potion in sacs snd tine is predetale. There test females at if they ate pase fe Source for vbich males compete, but femal ceice may steugy face {he fom of maleate compton, a for example, whee it fevors male comping together on display grounds (foe examplo, 8 Bea 1968) whieh fernales then search out (ee Below det "Fenale Chole"). Cervdsifer inthe extent 1 whlch faales ar clumped ln spe of rat omy dspered (deVos, Broky & Got 1987) as do antlops CEkonbery 1965), end these diereness carl in a pretcable way wits diferesce in mate stiibutes, Genel maleate aggrsson wil bs tho mote terre the greater the numberof fale wo mses ae ghing over ut ay hen omen, Searching behavior should be more important i highly elpeed specks expecially ifthe dspralicombined wih wnpreitaby ‘lumped in tme retro hlhly seasons! bretders in which many females become seualysvilable fr a shor period atthe seme moment (lors ample, explosive breecng fos; Brie 1965, Rivero & Bees 1960),whey digo Brees (in in) ar pein ae a chiar tess Van Lavi Ooodall 1968) in ih tas red ore ots ‘enon thought te yest Oe elt of exten clamping hte Slt mote il fo acy ow at be extent oz we he ie copula with on fol, nde of oor foals ae nl noouly bg nsenited Dera in nea let wien coun Wis - lone tad reptiles: the species ate dial to cbtrve aod fow bebavoal ae of any sor have ben recorded ex pone, however, tol thls core ‘sation between human lgsocaes nd speci in whi felee ne lager '6 not acidntal, Humans tend to be more Knowledgeable about thors species that ae also ative cumaly aed stonly dependeat ok sion, fr ‘rample, birds and lage mammals Tt may be tat male aggresive ‘nore strongly sete in visually created animals bros vslon proves ‘ong-ange Information op the Behr of competitor The male eat or example, eatly observe another mal pinning fo copulate and can een ‘eekly attempt to interven (or example, baboons, DeVore 1965 ané seg {rocte, Seat 1942) Marmals and birds nls tend towns low, Bed chtch sees und this ‘may favor relatively smaller fale, snc large female sae may be el ‘ively unmporant in sepraductive sees, Ta many fy Hed ond ae ‘nander female reproductive success measred by elitch tk i knows {9 conelate strongly within species wih az (Take, Wibur & Tiley 1970, ‘Tey 1968) ‘Measrng reproductive succes by frequensy of copuiaton, 1 hive ‘alyzed male aod female reproductive sues sty funtion le fn ‘oli garmant (Figures 79 and 7-10). Both sees show «sgiecet pon Sve correlation betnen sae and reproducive cee, ot the tend ‘males i slninoty stoner than the tend i feraes (p-<01). Cone stent wih his endeny, males grow faster a alles tha fle (Fig. ‘re 711) and each an adule weaht two td one-line hat dk feces. The ex rato of all aninale i unblaneed in fave of fermen, hich would seem to lndlate diferent metalty, bot the factor that fight produce the dierene ae not known. Malt are igly gressive and trois, aed large male defend covespondiogy lege tenors ‘win many esident fle. No data are avaliable om sas end sce in ‘eave encounters, but lathe cloely rested. (and beheviorly ee) fini), linatopus, BS pee com of 182 daputes shred In ie Bel ‘te won by te larger anil Rand 1967), Fetes ay oul eve ap at ‘le, but eel hat leger adult fora ay eps ighly more aceareal invert at Seva Selecion 18 ] i i | Piure 7.10. Reproduce sce In mae fale A gma 0 fe In a te Reprotuctive secs resid hy the nor of opuaon obtrvd per amber of inde or emai) in each nonoverlapping 3 mm ae exepry. Dut combed om ‘espa vie ot are rae tne 169 end than smaller ones, and this may partly be due to advastge in feng ‘trove agressive, ins lage fenales Wander sgn amily more widely than sual adit nee, At acne interpretation (based on ecological computlion between the exes) has been propose foc sexual dimorphism In size among animals (Selander 1965), ond the Interpaaton ey spy to nals (Schone 1967) Cenanly more is invaved in dierent nals mortally thin sa, eve in species In which mals grow to large si than fem, Although dale show convincingly that notional firs eonglyatfet han male se vival intro, a sexual diference in sn among hums ls aot tested ‘nl the twen-fourth week aur canseption wheres disses in mote tly appear at Soon a the twelthwesk, Stes eta (1980) have stows male rats exe four times the prc fereles do; the difereee removed by casation. Sice mer fer more tm prit-deteint Figwe 79, Mele end frale Ante gee coping face down fue ies than female (ley gin less weight and survive aswell) the vee 7 a ihe tak of cocoa rev, Pha By 3H inked proteinva, apparently ulated to es, may bea factor fo causing ‘ne lower male survival in Wd ate (Sehein 1980), (Phe eontecion betwee16 Doneee £ rvens igre 7.1, Male ad eal rth ae In Agel a furton on ti see baud an ammer 1970 rcapre of anima mar 3 (od mona boinc har Sean ited to ear of dat: {nate how mi ager aa hen iar age ele othr seul at prlcinucia and male cepodontiveaucess ft obtcue.) Again, although human male suvia smote adversely abeced by poor auton congi- tlooy than female survival, Harton (1948) presents eidece that the higher metabolic ate of the male i aa importa fecorInresing bis unorblty 0 reny dites which ske ales move avy thea fe- Ta. Likewise, Tabor Daswonn (1954) axe tat greater male mor taliy ln the dete, Odocoleus hemoras, resus from 1 higher meabolic fate High moto rte coud slate fo both aggremeness and searching behave 1 rpecdactiv uous nerense move cplly in one sex than the other 3 2 funtion of ago alone (fr eximpl, though age-dependent experince), thon one woaldexpet a postponement of serua warty in that vex and tgresier chine of survrng Groagh sual of se than in he oppo sex. "Ths the ad sex rato ight bo Saved i favor of the eal maturing Sex butte ex rai forall ages fae together should be blaed fn favor Of the later maturing tex, Of cour, H reproducve success for ne sex ‘cases strongly aa futon of experience and expeince only pty Perna Investment and Sut Slecton 65 ‘omslates witha, hen the sex ay Be wling to ue need mortality i thie mortliy Je svcendy eet by Incense In expec, Slander (1968) hae sopgpted tht the tendency of immature mae Blackbird t2 fxhbw some mire eharactry say be adepive fa Hatt Inremes the male expeene, although ic algo presumably crates is sak of octal. Data trom mammal (eviewed by Biunborg 1965 and Brown 1966) and from some salamanders (Mason & Shoop 1970) and momeroot ards (inkl 1967 nd Bint 1960) sore hat males often cocupy Ieper home anges end wander more widely tan females even when males are aller (Gla 1965). Parker (1970s) har quaiied the Impocaace of reolly td earhing tohavior a dang fies Tf females are » ispesed reours, then mle moby may beers In exposing th male ta lrg number ‘lala emle.Agsi, males may be wing to eur peter mealiy UEehiseucondy fet by Inteats In reproductive sues. Ts fete hou only allot the rule dig te bretig sean (Kichaws 1968) ‘ils faeore relevant fo moby (each at epee, opi or knowledge of the environment) eed fo be developed prior fo the repro sesso Lindburg (1969) hs shown tha macaque mals, bt Bt femal, ange ‘woops more frequeily during the reproductive seszon than otherwise and Thats mobil nereves mle reproductive sce at eared y ter ienay of copatton, suggesting at at Teast in this spec, eater ‘nobly canbe canBned to tha repeodstve season (se alo Mller 1958), (On the other hand, Taber & Dassans (1954) presat erence that os arty st month of age ral er wander more widely trom thee moth tis tan fomder~a diference whose fonction, ofcourse, ot kaowa Shuler very early erences in mobility have been denoostted for «| Taard (Bla 1960) and for sve peas, lnloding man (Jensen, Bobbi & Gordes 1968), Female Choe Althoagh Darwin (1871) thought female chaice an Inpertant e¥lar Vioary foresy mnt weer se him have clogated Itt tial role (Huney 1938, Lack 1968; bat res Faber 1988, tad Ozians 1969). With ‘otble exception the ety of female cole ha ined Ise to showing {hat foals ao sleced to decide wheter pote pater fof the Fiaht spec ofthe ight sok and sxualy mrt Wate the edaplie ‘ale of such colts obvios, the adaptive value of saber dierimina Tins among broadly appropie mle ir uch more leat fo visualize fr document, One sede Sot theorstin argumesss fr the edapie valoe16 ewer, aan Of such female eee and deed data on ow females choose, Net of tea cits met By tase who eal asebe to female (or mal) hole the evolution of such was an the rltve haloes of both ie man sexes (Hersh 1968) ofthe large sis of hom female rest (Mons 1967). 1 review here thereel considerations of how female rght be expeced to choos among the avele mals, lag with rome fats on how females do choos The efets of female choice will depend on the way femsie chose. If some fraes exes » preference or one typeof male (genotype) le ‘thers mateo andom thon oer tings belng equ seleston wt epi favor the prfowed ‘male ype aad the feos vith the preference (©Dosald 1962). It cach female hs a specie image of the mle with ‘whom she prefers to ite and if thee i decreasing probably af fe ‘le matog wih sale ats funtion of hs increasing deviation tom het Droterred image, then iis via fo show that selection wil favor di Hons of female prefrenoes end mle atubuter that coincide. Female hice can genre continooss male change only i females choot by # ‘eave ruse cham an abslute eteson. Tau if heels tendency for feral to sample the ale dstbation ad to preter one eteme (C0 ec ample the mare bright cloed males, then elesion wl tone the al dintibwon toward the favored exteme, After 8 one gineslon Ing, th ‘tibution of female preteens wil tleo move towerd 4 gear per ‘centage of females wih exteme desires, beenze the gandupiters of f= malt prefriog the favored exteme wil be tore nureras than the sracdenoghtes of females favoring other mal stb, Unit coon ‘ling ezedonlnervenes, hs female preference wil Sat pont out by Fer (1958), move both male abate and fale peernces with ineretsing rapidly inthe sams deton. Tae fale preerence fe eapable of overcoming sos countervailing tlestion on the mal ably fo ute ‘ive to reproduce, Ifthe inresedreprocotive soccess of the favored ‘uals when mature oats his ehances of uring to reproduc, “Treat atleast two conditions undee which one might expect females to have been elected fo preter the exteme male of enmple. When two ‘peck, realy spesated come tpster, selection eapidly favors felt ‘Who ca dsciniate the two species of alae. Thi seleton may favor fe Tnles who prefer the appropee exueme of an avaible sample since such a mechani Would mininine ming mistakes. “The rata section of females with sush a mechenm of cholce would then ile ssa Selcion in the same drei, which ia Ge abence of countering Selection Would move the two male phenotypes farther spar than neers fey aveid mating ear Darel Inesinen ant Seal Slecton 1 Netorsl selection wil akon favor female ably to dsiminae ale senval competence, apd the efest way too this ito tke the exten of fanmple, which would led to runaway sletion for male espa. This ‘ose lssaed in or dtl belo. ‘A in othr aspect of sexu! elation the dace of me investment nthe ‘spring fe faporant and should afet the cetera of female chle. ‘Waese the male laverte ite or nothing beyond le sexed, the fale soi to desde whch male oes te at genet materi for her of Spring, eeunng that ti ewig an epable of oloig i. This que thn etn be brakes dawn to that of which genes wil promate the suvial ‘other efpring aad whl wl lead to reprosutve sce assuring the sfpring survive to althood, Impl im thee questions may e the ral tion between her goose and those cher mite: do they complement ech tet "Where te male inves parentl ear, female chcice ray sil involve te ove guetons ofthe male's genes contin but eho lio invele, feria primarily Inve, qaestons of tho male's willingness snd ability fo bers good parent Wal be inves inthe offspring? wig, dove bo Ihave the niyo contribute mh? Ags, narl selection may for f= rale atemverets t complementary: do the male's parental abies complement ber owe? Can the two. pants work together smth? ‘Where males invert conterbie parental eae, mst of te sae cone ‘lone that apply fo female choles alo apply to male choke. The aerate ‘era for female choice are remmaried i Table 7. ‘Ben in mile sete for rp, peated copulation te billy to d0 20 Ie ot unlined, Ate thes or four sucetve ehcuations, for example, the concentration of spermatozoa le sry lov in some tale clckeas (Packer, MeKerse & Kemputer 1940), yet males may copulate a offen as 40 anes in an hour (Gobl 1951). Lew, sperm is eompletay depleted in male Drosophila melanogaster ater theft conseative ming onthe sine dey (Demeree& Krsfmenn 1941, Kaufmann & Demeree 1942), De- ratlon of copulation et in hal by the thi copaation of» male dang fy fon the anne day snd durason of copulation probably corltes with {em rnsered (Parker 19 70a) [a sane species females may beable to jndge whether addtional sperm are needed (or example, house fess Riemann, Moen & Thorson 1967) ox whether a copulation sa ss behave toaly stool (for oxen, sea Hons Peterson & Bacholonew 1967) bt in eany apse females may gunanie reproductive soeess by ma168 owen rvins Table 7.1 Theoret! etna for fale coe of males “Tics ot ejection male port lean Gy area esi ) coeatce ) ame (0 erly ono Guay ef eee Go ety tpn ore (2) eral sof pene (5), Stiomensnty cso ©. aay of mt ne Cy ng mi vet 8 Ing wih those males who ae most vigorous in courtship, since this vigor ‘may comelate wth an adequate supply of epeem and willingness to teaser ‘When the male completely depleted these eno sdvantnge in hs cope lating but selection against the mate doing so should be mich Wen han seleion aginst the fale who accept im, At intermediate sperm levels the male may guin something from copslaton, bat the fae Shontdagae be sled to avid hie, Slce tere site advantage to the ‘mao in concaing low reredstive powers, corcelation betwee vigor courtship and spe level woald not be surprising. Femles woud thee ‘be selsted fo be aroused by vigorous courtship. If secondary erucures sed in display, suchas bright Teather, heighten the appenance of vigor ‘uate, then selection may rally scantuate ech erates, Leonel, the male who has been sexy met suscenfal may not be eal toa with if this soccess as tepocariy depleted hi apann euply. Males shoud at only bo slated to reaver rpily fom copulate bat give convincing evidence thet they Have recovered, Tt not abu to eipgose ‘hatin some highly promiscuous species the most atoctive mals my be those who, having sready Boon oberyed fo mee wth several female, ae sill capable of vigorous diplay toward a feria in the prootsy of ‘hooting Maynard Smith (1956) hs presented evdeac tha, lve chic, female Drovophila subobscura aueininate gana lbced aes of tha species ‘nd tot thie Behavior i adapt! femsle who do not so locos leave about 4 as many vtble faring as those Who do. Females may Pare Inenant and Stu Section 16 choose onthe basis of courtship beavor: bred males are apparety o> tie to prtoom a step ofthe typzal courtship a apd ws outbred als, ‘The work ig purty interesing In revealing at deals of cour beaver may reveal genetic ti, such being inbred, but sues from an ariflealty. inbred males prodae soul lable opens, "hen, even in the nbonoe of female dierimiaton,oge Would expect very few, any labred alo tobe ava fa the alt popuition. Only Bee ‘aus such males were artlly elacted were Chere large nusbers 0 ase to females fa choles expedinents, Hd that selection continued one entrain fra, frase who chove inbred males weal Rave ese the ‘oe foals, Mayaard She study highligh the problem of analy the potent for survival of one's partners gece: ope Know of the elt miles ne mets hat thy hive sried to aalhood; by wht celia does ope do= fide ‘who hat fuvved beter? Tete female eon juoge age ten all other things being qual, She abou choose older mals, a toy have demos- seated thir copacy for lng suri A ober tioge oy nat be egal, however, I ed ge corte wth lowered reproduce sets, at does in some ungulates (Free 1968) through redoeed bly to inpregate the formle can judge the physical condition of male abe encounters, then she can icine gsi wodeenourished or sickly inclidal, fine they wil be wns to survive long, but discretion spas sch Individuals may occur for other reasons, soch athe presumed lowered iy of such males to improgmte succestolly due > the weakened conton, Th some very rested ways It may be possible to second-guess the future ation of tural sletion. For example, stbliig slocion ot been demonststed to bea common form of naturel selection (sce Mayr 1963) and under ths fom of secon foals ony be sected fo exces Unie ove dleriiaation against extreme types, thereby augmenting the tMfeets ofan sallang slot that fas osurted pee to repeodain, Mason (1969) his desensvaed tht females of he Celforla Oak Math ‘lariat ego miler extee in some tal, but 0 oot has ehowe Indepeadeat stabil seleion for the same alt Disedinaton galt extreme types may run cous to section for der; ho po Sle ole of female cots In increasing or dereasing vey i dacosed Below form of omplementary Reproducive sucess, Indepeadent of wii to survive i ease for the female to gauge Beease she con dicey obsorve dfereaces Ia repro tive success before se chess. Asking feature of date on ek bebwvor 1 irdeto tendeny for females fo chooes males Who, trough corp tion wih other mele, have already increased tei Ukethoed of mating. male cole thea geal augment the eects of male-male competion. (On ths lk grounds thas Sean obvious reason why th may be scape.170 poner evans ‘By mating withthe moat dominant mle female cun wally mate more ‘uly, and hence me sully, thn she chocats les drat ini: al whee tempts at mating fen ret In intereenee trom more dom oun males, Set (1942) ha shown tht my raigs with ss dominant fndvideals occur pally when the mare dominant invidvels are un 25, eltorbeease of text existon os Tong wig line, to uk Servic the female, Likewise, Rebel (1970) has shown tht dominant fo ‘ale prevents esi dominant nde tom mating vn she has mate, ‘resurubly to shorten her ety abd fo copulate whe the dominant ale Stil con. A second reson why shooting tomate with more Zola oles may be adaptve is tht the female alles hr penes wih those of & male Who by hs aily to dominate other rales, be demonttaed Hs repr use capac. Iti common observation in cere tht females lacidly avait the utcome of male slle to go With the victor. DeVore (1365) has quant the inpotnce of dominance in mae baboon ss) ces, emplsizig the igh tegueney of ftererence by othe mules fopstion and the tendeny foe female choice, when It appre, tbe ‘oresed in favor of dominant mule. That provogssicoee may Ines ‘heal wih whch les court frase suggested by work onthe back rouse (Kult, Bossa and deVos, in pest), and females muy preter Innes kis a courting inact Beco hk bil covets wih previous "Tn many species te ably ofthe mal o find reese females quickly may be more important than any sili 0 donate oe mals Ie 3 io, then feo colce may be coniersbly implied Ihe ft ‘ale to Teach her exabishes thereby» prima face eae for his reproduce bls In dong is, in which faraes mus ate quickly whe the Cons 1s tes, male courtship behevic i vtally noneient (Parke: 19708) “The mele whe fist laps on tp ofa cemyarved female copletes ith tt This lace of female cbeze may also eel som the prin fete case the frst mal exablis for his sound reproductive abies, Such meche tank of cholee may of cour confit with other erteia equleg smiling ofthe male popustion, but In some species thi semping could becarte out prior a becoming orl oseptve. ‘There are goed dats supporting the mpactanee of complementary asses to fermi choke, Atcortatve mating inthe wld as been det cate for several bie species (Coch &Beusire 1959, Dont 1989) ‘nd digssonatve mating for» bird specles and a moth speies (Lowther 1961, Sheppard 1952). Pat & Eheman (1968) have demonstrated the tendency in several Drosophila species for females to prefer ating withthe fare (pein choles experiment a teadeny which inthe wild lads to 0 form of complementary, since the ferle is pressmably ually the common type. Thess sties can all be explained play i tems of ‘election (or gresteror ser gees dives, the frie chosag # mile Parental Invern and SesutlSlecton m whee genes complement hor ove, producing an “optimal divert inthe offspring Where male paces cre i involved fales certainly sometines choose rules on the bs of thelr ability 0 contbute pana care, Ocans (1963) for example, has recently reviewed argments and data soetng that palyeyay evolves In binds when booming the ssond mae can I. reedy mated mle provides female wih eternal pretal contrib: ‘ion than Becoming the st ale ofan vamated ral would. Tis wl Bo $0, for exumple, the arendy mated mle dslende a terstory consierly {peor to the omated males. Variliy In eony quali ertany 0 us ln most terra specie, even Inthe ia whieh terites ste ot ied for feeding, Tnbergen (1967, for exaaps, hes documented the tendency for cota eos Inthe Mack-haded ul tobe les vl thie t predation If femles compete among thems for rile wih food terroir It ales extest stole sa wel, then fea cole for veal abies wil again eto aupmeatisra-male competion fr the Felevant resources (such a ears), THe ros abou form of ths 9 lection ithe nabilyof entrar boling mele toc fae Female chlee my play # roe in selecting fr increased male prea, lnvesent In the vondenaer, for exampi, food ought byrne sees to nt on him as eb aphvoite: he roe to feral aed coats berth the food, soxgesng tht the femal would nt usiy male hou ech 4 pit (Calder 1967). Male aril sare invested lfc copation pre Somably nota esl of female choice after copalation, since she no loner br acything to tapsin wth In mat bids, however, sales delend tt toe whi nia the females (Lack 1940), Shee males withoot ‘sable terol are sable to attract 2 mate, female coi may play 2 rola a intaning male torial bebavon, Once smal ha loved In ‘tecory inorder lo atract mate bis opine ater coplating ith her ray be severly Inte. Diving ts feral ot of ie trary would a= tos cetaily eesti the fs of hie invesement up ust thes, He could Cash another territory, and in smo specie sone sunle do th (von Haneman 1951), but in mocy ape this may be ef, Isving Rn wih he opton of ning. more orl, the female be bas elzeady mat Female chic, then, exec’ Before copition, may Incest foie the ‘ale oer hi parental investment cer eopulstion "Thee no reason fo soppore that males do not cotpete wih each other to par with thee fmaer whore breeding potential appears tobe High. Darwin (1871) asgnd that fommles within speck brsding eal fot rongenlc reson (sich ns Being In exellent pysioe conden) would produce more odspring tan ltt breeders. Seal selection, ho ergued,woul favor ie competing with each ater to pair with such fies. Fisher (1958) has nlely sumauriaed thls argument, but Lack (1968, 157) dismiss tas being ot very cogent ince "te ate of Breeding in bids fas beeoevlvedimally i relaion to tWo eileen factor, ‘namely the fod supply forthe young sol he capecty of the lente 10 form eggs." Thess fas are, of course ally conse with Dain sre ‘ment, sine Darwin merely soppotng 4 darlopmental plik that Slows ferates to bred eve thy ae capable of forming the eh, and aia preseatedelshers ln Lack (1968) support the argument that fe ‘nals breding ele for aoagensle reason (wach ar age or duration of re bond) are more sacesfl than those Bending ltr (ee, for ex "imple Fisher 1965, and Covsoo 1966). Goforth & Basket (1971) have re- éeoty show tat damian males in a penned Mouenng Dove popuaton preferelly pal with dominant females; such pe breed cation and rode more surviving young thas les dominant pis t wool be inter fsting to have dtled data tom other species on the extent to which rales do compete for feoales wits Higher breeding potenti Mater are Cerny cen nly aggresive to females Intuding in that tere, tnd thi aggressiveness may act as a sev, adntig only thee females ‘whore high metintion correlates thea gg laying and high seprodoe te potent Thre good evidence tht Ametican women fn to Matty 1p th seciossnomi sel, and pies aretventee ding adolescence fetes such movement (Elder 1968). Uni reseiyaieh 9 bie inf imale choke presumaby coveted wih increased reproducve size ‘at the vale, any, of farsa Bony for male reprodetvesosess i car, ‘The importance of choice by both female ad sal fora te who will ‘ot desert nar priate nek oti the ple bood has Bae enphaied 1 an eavier secon ("Deseron and ciskclry"). ‘The, knprtanee of complements x docomented nx study by Conlon (1968). In many species male-male competion combined with te imparance of seme rsouree tn theory unrated to tals, soch as ovipnion sites "Enywiltgute agaist female cle for mle chaste. Inthe deegonty Parthomis tener als eoropets wih eich other lo eantal ecole eon Inning good oviposition ses, probably because such res me © presi able pee at which tend renpive females ané becure sper compet tion in nsec usualy favors te let male to eopulte prior to ovpcrion (Parker 19706), ts cear that te females choose the ovpotion site and not the male (scabs 1955), and mal oui geared to advert good viosion sites. A mle maintllg stetory containing 2 ood ovpoe Paral texans and Seal Slecton m on sn ty cotibtig pre ives vs ht ale ‘tue ea he aoa! : coe roto smb am cay npr oe lt of al someon hs spec hokage ae ders Sowing sta ferteton Sa em alesse a depued ‘hee spc tote eon of mle pa eee Fetal tle for got ocr ste wold end oe ay as sine! a impoig tn and anced ea eee tel ele wo a apn cag oro {Rb sony. h'srengeat won sed ef eae a fos tod deveopaen mite apna See es areal ct army snc el she ces Won ae ae So faor mal couralp fed, wish hhvons Need ea ‘al ol chose my ao ner mas ao ae tay ea Psion tsi doing eed ho pay ee her females dunped insu th eect may soe fle coe ume impose ine meney aoe a oa Cre fo donee rl may pve en fel cng fe Ba te abby ren up wad aad by more Sanna Soe Psat ene ceca nthe ts atone apes A Victor outlined above, a eae eee Suinary “Te eave pret iovesinet af the sexes in hei ou i th Key wile cntoing the operon of een sceton, We oe ‘es onieabiy more tn he Ob, embetter wet ones song hemsle o mate vith mombrs of e omes, Whee cerns i oq, sexi eiccon eho! operat inary onthe tos See ter of rave pare ives in apd toy eco shai oe fence by te ely eveutonarydilerenten into bie se coe ea ig immo, and sexual seesion nso te pate ey fare ivesineat. ‘The tine sequence of petal mening any by sx an Important parameter aecng sec In ike bok Sea ek yest conicable puental cate the ininiue nil neato Casualy the fen) is verte to Seseton, On te ce neaae ‘species with interoal ferilztion and strong. male parenlal investncat, the mae is aaye alae to cckldys ack anes nes to the evolution of adopalons to decree the isenoly ay Nd Countrcapttos Females analyser hiphee mati rts than mls in onogemoue ‘ids, bt in ponmonogamoce bis and all her psp na tetn oeeny Tene suffer hier rte. The chromoromal hypoheris uneble fo account fr {he den Instad, an adaptive ierpretaton can be advanced base on the Tele parental ivesimet ofthe sees Tn species with Itleor no male forcial investment elation welly Savors mal adaptaons tat Tad to High reproductive sic gone or move breeding seasons athe costo tread moti, Male competion In rsh species can oly be anal {deal wien fe dtdbuton of females in spice ad tie Is ropely feserbed. Duta from feld nies eves tht ia some spect, sz Imoblty,expedence sed msbolie ate are pont to ae reprodve the sie "Female choice cn augment or oppose morality sleaion. Fencle ‘hole can orl Ten to una change in male morpology when females ‘hoot by a rltive aorta sbsolut standard, abi podobly some Times adaptive for fonels to ao chose The rlatve parcial levstnent ff te sacs ect the ria of female chece (ao of male eho). “Throughout, I empbasin tat sexual sulccion favors diferent ale and female epraduetive srlegies apd that even whan exteaibly cooperating inet ak ele and ferent are aly eae Revenences Barton, G. A, 1970 A medel for the oven of piped pole. “Eelaon i 54659 aioe, M. 196, aarp: A elles wud, Chicago: Ai. Brew nas wast Drosnpita Herdy 2349-368, ‘Beshe W 1525 Th warigetesTiemau Cpr area (Gina), Zoionen 6: ISEB, Belg Re i B Pond £ CF MooLend 988. Sx rts of some Mione- ein olan dneion Midland Nouri 9 82324 ovine, 65) Hal 1939. Avvo the span and poral ates ‘fat notin and threo fo erat ad eer Pysiogeal trace {hin Inthe ifezyon of anima, 02 G. Weertcmne & M. O'Conner, ip ett ones keh ft [Bit The hay cada: Unies of Tes. oulry, 2 Fy Verse, 1960. nouctan ecto der ongles de "Pare Natal Aber rl: Ian eg are Nationa do Cong Belge Bragg ALN 968 homer of he mii, Poles: Unive of Pes rows £1956, Home range aed mera of el mama, Symp ‘ltt Zaoane!Sooty of Landon 18 111-10 ‘eat 1960 A took 6 pregeaney Inthe mau cased by We proxi of "arange len, Jaret of Repreuction and Feri 1: 96-103 rutny EM 186d hing rogeeney inet! poultios ot pcs ud but tet deerined Sy spemaiophert coum Proseding ofthe Naina ‘Academy of Scece GLASER. coer A136 Breating bobo ofthe Rndruster, Gesece caf mt, dik 04 957-538 carpi, ©1967 Atgesog 2 ail strc ia avai is a Lord “olay si. Mitad: Coombs, Ma Unies of Miso. Paral Invesmant and Seva Steson ns Chapman A By pg, A.B LB. Cul A. Cote, 1998 Sex ts of el cae. Pro casio American Soy of dni Proncion isp 508 eto: Ba Sow ome capi Rane (88 POSE Ingo Cue? Langa W Moore 1B. Drea, Linde ting "esni onal of ia Eetoy 39 2St Se OE Hog arias wl 208 ie tence of ep bond aad age on he edna Wo gf he Kivake gll Rise wae. Jowrel of nina Bevogy 33" 268. ‘Con, 1950, Some via satis of i pe on cracked mou alg [ti Sox at ad mor nth ee a ae Darvin, ©1671. Phe deca of man, MEG TL The deen of mam nde elon Ldn Demers, Ms & Kautinnn P1941, Te regaled for Dro paint i up of mitre parm: Amat Nanrla ese aTae ee 1965 Mile domes and sing thot i boon ao eli ce. Eran Beach New You Jefe Wity a Sones eon AvP Bro, &V. Oi 96 A encyclo ofthe North ‘Exton, T. H. 1941, Notes on the lf cra let Reve 3a wien # mo mlz of Wife Managemen 9-98," Corsa Qt xyes, F970, The poor of inet reproduction, Oxfords Pegnon te XJ, ohne Vettes a Semin, Sang ‘adie wn Saperdtien(Sygnahiuse), Zerach, Mess Vso a ihr 1969, Tag and caving in the Lays Alb meat kom 08, Died i SRS SH Pet ity of na ato. Nw Yok Dore MEA E196 Radu beer snp London snd New Yr (Cai M&W. HE Mower, 1970 Life hori comeguense of ur a (ati 1971 Fc feng pemitiv and ‘ntng remit tx. i Suan i the tctology 8 0a) boat of ung, ated BH. Kew “one forsee ones So an, wont Ws °T, Bnet 197. Soe gen opened Mouang 1, 88-9, Tiny of Dendrbaer attr Copia 8 chy. American Soe16 owen 2 tens "Sense 30:68. . st on ep te cries camsacees esto et WLS 0, ore Sree nthe oi “rc antiga 1B, 8°, Hamon 8 , 2. Mesor, 196. Daron of ie and CEE ee ania wats mt me OGr Ektrtnry sx al. Suece 156: 477-48 sa oe Eee Ee «marten ct VES a Go a tg of os a age bi trees cara Saige WE Sa « en tone ne we cn re os eg ne eek wee tl aie te Shae fen Roemer tae Pas "Stonotngestsand pdemar fate worl oa fA eet vite f 1965, Behaviour ofthe Back grows during the sting dla. Parcel rowsment and Sexual Secon i” Kopin F194, ue Sessler fet en 181 Bourn, 8.61.06 P,P unig cao sale in Black Grouse, 15:h Cong. Inerm, Orith, Toe Wagan, Hd. Spe BD Moe 6 Sl eto Se CE Be Oman ack Bisel Se, Cone a 265-246 Gare Sie etd ep fen ner New Yorks Oars Fm ht eee adnan for ren ds Lato: Meta. Jet b 8R3 Foon 10) Sacel Sa ee Seco 1B Gta Sara ae SB hele Seiya er i aD) Hrmerp Spee shee 3 ses ome th 619 Rc mck: ma et me lr Leni 2 1 Pamartien ee winced aon, Zoe aiblcolis (Gmelin). Canadian Fownal of Zoology 39: 28-29%" le WG Ba. 8h ee See tee Eaten, Wu ist crc Ea Maghon BM & Stop. R970 Homing teva, vente, sd tome gt linac da ea Sk a ih 6196 Vn santa oe Coin CoE od nt Droptidoe). Evcluien 2: 35-0 eased age Sin TSS Ret ming beter apd sleet a Brovopiasuboiscrs Journal of Genetics 3a. B29, , ens 1059 The a wt dr ir 2: 16-19, Malsinces Sena. Ze ce inate A American Naturalist 103: 589-604, * “lm mai on ono Joe The reponatv hao and th ature of an slton Tae ect them Fe le eee eras tm Pal Cy 8 Ebene, 1969, Seca sleton in Drovopil. Yo Evolonary tigger he Teed ic RTE ne EE ee mmr ET a mn em ce Seiko ean we aes in Sef a etn ec a ul SEES eran cone ono Sent dae i Ss at annie ah ae meee ae See ooo CSETEE Matheney 9:07 rental Tovesment and See Seen 1 Sane, RK. 1965 On mtg pet ad sex! dB 1963 ” 5 sleton. Arca Note P66 Sal dna se itt nde wo ai ‘Condor 68: 113-151. " bi ie Bede se (CN Se, Marin 4M, 19 9 tn oily Oe Anan eo Psy 1 ea soln ME Asa ern ee ee dominila, (Le) Heredity 6: 239-241. wh Povae Sur 8 Sea eo oe Wi Rab Ore ans cuniculi 1) in We Was, Proceeding ofthe Zoaones tock of London 122: 417-434 “ " coma she Wine 97, Comp fn in pinta rd, ak Seay 36. Sol nmin Henan gu, Sc om shan amon path 5 Amant hye Unley os F, Dauman, 1956 Ao diteense la vce a yong clog lake ee Yuma of Whe Mange B23 spl ttt Demosnltne laments Byohin 2 agate EM te, N- 1935, Fl obra of Eat Creel |The bebe ‘fhe Kesinected Phalarope (Phlopus lous) Spay ane 2 pig Bralton 24: 38-16 ny sme nant Bn dea a ae se wn Elk. 1963 Sel ttn, srl, 8d ml wemerioen Naturals 91: 405-406 “et Resumes