_ovruine, Ways of laking a animal behavior Clnsicl ethology and innate Bohvios | Communication in bows 1. Bed action patterns in vercbrates Other functions of tate, strctype behavior 5, Buran elnlogy 1, Behavior genetics ‘A The ccs of behavion—naure vs, tne 8. Hentai W, Learning ‘A Habituation Rese lening 1 Cast ending, 2. Operant conditioning , Caters loring tng learn, Imprinting VC Migration VL. Soe behavior communication 1. Modes of communication Bomninance erarchles Tortoralty Sevual behavior and sepreduction 1 Pale bonds 2. Cane ofthe young, Pay Soobilogy E 52 Animal Behavior “LEARNING ONJECTIVES _ [Alter you Ive read this chapler you shoud beable to 1. plan how a imple behavior, such as rine marking In dog, canbe interpre frm the pot ew of (a) casual ethology, (blaring, (obehavor gent Is, (@ behavior ology and (6) meray 2, Disingush beer posiate and lms foeors 38 ‘uses of anal behave 4 Dinu fe action pater rom reflexes and ol ‘onal Behaviors 4. Gio examples of how displays may have evolved through the aualization of ese complex behaviors interpreting human behavior in Tern nate be Favors in animale, redirected Bean, displace ‘pent behao) 6, Diss the Infisnce of genetic and lard compo ‘ents on behavior i the stan individual and in pop Tans and explain how the cnenpt of Reta x ‘een the ter we 7, Clay lene behaviors you encounter in animate at ‘samples of) css conditioning) operant or ‘Song. ()senstzation, (Q)habtuabon, oF ( Sight ening 8, Docusr the apie sigaitcnce of imprinting 5. Disaet how tel ane and handling te ary when nimals forage optimally im good ond poe habia 10, Postalte toll advantages for migration. 1, Present the concept ofa dominance erry, ving at lest one empl, and speculate is possible general ‘apuive sigan and socal fenton. 12, Distnguit between home range and teary, and [Bre time theo aboot the adaptive sgnance of 13, Banus theeaptive value of couteip Behavior and dese a par bond 14, Summa the emphases of soobiology A tamiliar canine behavior is urinating on a lamppost. Why do dogs do this? The anewer depends onthe level of explanation we have in ind. Anima behavior, succinctly defined a5 "the movements of animals 12331234 Chapter 52 if sneer to smo” das wth sv els of ens hy an remade the physiological mechanisms that cause the behavoy ay level net volutinary processes that account fri origin and develop wl a eer ihe mr apc oe animal ea ei dating ad organizing thi onan Ee — WAYS OF LOOKING AT ANIMAL BEHAVIOR ‘To an ethologist the question “Why do dogs do this?” might meen “How Te ee Dekavion, called rine marking, help a dog, survive in is natal sephatt Broadly defined, ethology is the comparative study of behav. Fahettgy fear on ini (inherited, inbors), specesypial Beira cee ae ature and on how these behaviors adapt animals t ther enion- seer polos also ty #0 piece together the evolutionary history of tune, oo the question also ight mean to them, “Why di dogs evahe ae ng aa than another behavior?” of, “Did urine marking rae a preexisting behavior, and if so, what was that bhava’ Thay imake the reasonable esupion tat sanate behaviors Hk hs ne sey rs veoponsve toatl selection (Chaplet 17) ate estomiad features ‘ltvior geneticists are concerned with the relationships between genital bebavion. Accordingly, they ask the same fundamental question Bees netics ack about biological characters: What are the manner and ror hich harecters arene? After establishing how ss it Thgleancegencially among indivicual dogs, they might analyz fing (Re eee matings to determine the genetic mechanism that ste phase inbeitance ofthis tra ‘iperimental o comparative psychologist, particu those wong in ta tedton ofthe brio (eg. B.F- Skinner and his colle, Fe pnp on lara bchavies, hat, behaviors acquit or ta fat by preous experiences. Their objective iso predict Eehavor nan fe evince wth the god of contoling particular behaviors under Tac ceumstanes. To tis en they determine experimen the BRS of apecie vaiales—such as dit, age, ambient temperature {he presency af ther animals—on behavior. Even insted beavis Ne arr ing can be modified to some extent by environmen arate, Pane ple’ dog, after being punished for marking postu FeAl ileal at spot but no actualy urinate, even when unten by its owner Most experimental psychologists carefully contro he tions under which they cay out thelr expesiments. Typ! ‘domestic or laboratory animals a subject rs eertonyfocuzes on the evlaton of adaptive bavi relsten to the envionment. This discipline uilizes theories 204 ‘gues fom both classical ethology and ecology. Wheres dase ‘ule suay concenteate on indus o smal groups, beni B15 genrelly sty the dmamics of ebavins in eli Basins particular attention to behavioral fferences betwee, paying Paice between populations of dierent spe Ay aaa aac is to quantify the behaviors of individual 92 TOT ie ra fr oyporate the data into a theoretical model 1 EP a, apt sea to ts envronmest In dealing ih 50 ar peptone, behavioral ecology often relies heavily on athens oratory on thy oH in ” he Pp ho ji | a Ms e i[Neurobiology secks to explain the mechanisms of behavior in terms of the structure and function ofthe nervous system, To return fo the example urine marking in dogs, a neurobiologst might bepin a study ofthis {ehavior by identifying the specific parts ofthe brain concerned with i ‘tivation, organization, initiation, coordination, and execution; the neu ‘biologist later might determine the biochemical teactions in these brain forts that accompany the behavior, Newrosthology is somewhat narrower [Rscope than neurobiology; it concentrates on neural mechanisms that Underie species typical behaviors ( observed in nate), that, the be vows traditionally studied by ethologiss, "These areas, all dealing with how and why an animal behaves ast does, ne generally regarded as subdisipines of animal behavior; but many of these areas Ovetap considerably in subject matter and experimental tech riques, and most have been defined in more than one vay. The term “atology,”" in patcular, has been variously defined, Some ofthe subdicciplines differ more fundamentally than in level of planation Because they ask different types of questions (nd thus require ferent types of answers). Questions about evolution arebasially histor ‘al questions that must be answered with educated guesses about what Ingh have vecurred during the evolutionary history of an animal. These long-term evolutionary causes of behavior—called ultimate factors ‘Should be explainable in terms of experimentally verilable mechanisms Such as genetic change and natural selection. However, the hypothetical ‘xepunt of what occured is itself not verifiable Ta contast questions dealing with neurobiology, behavior genetics, and “experimental paychology are concerned with the Immediate conditions oF tnechanism-—the proximate factors—that couse particular behaviors These questions must be answered either with facts that are verifiable ox- Pimentally. or with hypatheson that af last im pincphe ae testable CLASSICAL ETHOLOGY AND INNATE BEHAVIORS, Most of the frst important contbutions to our understanding of innate ‘shavior wete made in the 1930s and 1940s by several European etholo- {ls Three ofthese ethologits—Konzad Lorenz, Nikoleas Tinbergen and Kael von Frischreceived the Nobel Prize in 1973 for their work “The behaviors that attracted the attention of early ethologists were those tit pique the curiosity of many sensitive observers of the behavior of animals in nature. Yet pechaps these extrordinarlly creative scientists thought somewhat more deeply about what they saw. Also, they were sscptonally ingeniows and crea in desing experiments to tet thelr Iypotheses Communication in Bees ‘innate behavior in invertebrates that was studied extensively by von Trach and othereis the "dance" of honey bees Figure 52-1-This intiate ‘etavir allows bes to communis the eatin of food source to ather bes in the hive. In one species a worker, after loaing ‘distant fod sure ets othe hive and perforin the dance ona verte surface of th comb Ie moves ina igure 8 the wo loops sharing "aght en” sing the stright un the bee waggle ls abomen These movemen's ‘muni the ection and distance of he food source from the hive hey are responses fo environmental ai that are rather sue by nan stands: gravitational pl and the drction ofthe sun. The de ‘ston a the food source, restive to the sun inde By the ange of Animal Rebar 12351236 Chapter 52 FIGURE $2-1 Dance ofthe honey fre indicating dieton ane ie {ce of fod source In relation to tenis * \ Q roa ome the straight run with respect to the axis tha is perpendicular to the gravity tional frce ofthe earth. The distance ofthe food source fom the hives Indiated by the frequency ofthe waggles and the speed of the dane ‘Cir worker bees follow inthe dance then leave the hive and fy the comrect diction and distance to the food source. It is wsualy argues tat this behavior i adequately explained by neurophysiologcl processes n+ totally unecore insect, Hovrever, some scientists have maintained So atten should not be excluded asa factor inthis behavior. Fixed Action Patterns in Vertebrates ‘A dassic eample of innate behavior in vertebrates i gg-raling in We foul, When an egg is removed from the nest ofan incubating 65° Dlsced few inche in front of her, she wil reach outwith her neck an Pi the egg back into the nest (Figure 52-2). While peshaps apps" Fither nsigrfiant to the caval observer, this behavior suggest 2" ber of important ethological concepts to Lorenz and Tinbergen, wh fully studied its characteristics inthe European greylg goose ‘After it was conilmed as innate by observing itin inexperience cqgeoling was analyzed in terms of i behavioral componen’s ‘led that while the goose was rolling the egg back toward the ‘ge veered off to the side, the goose steered it back toward the #8. MP {hue component, which seers the behavior relative tothe enon, tistingulahes eggvling from simple reflexes like the knee jh TA (Chapter 6), tn typical reflexes a muscle contracts involantanly we" se est tthe is = ——atteP ofthe body is stimulated, but there no steering component. In the case of the knee reflex, the lower leg extends in response to a tap onthe tendon ‘egardless of what stands in its path. iteggrolingis nota rele, then we may ask whether s the same sor, of behavior a golfer exhibits when he or she reaches out with a olf ub to feeve a ball that fs rolled into a ditch, Human behaviors ofthis type vito ehaoiors—are based on conscious choice or decision. Although the mental state of animals are not dvelly accessible tous, one compo rent of eggcrolling behavior sets it apart from most human vaitanal be- favor. I the egg is snatched away from the goose while she is erievingit she continues the retrieving movement con though here i lnger an eg 0 rere, Thus, once begun, the behavior Is rlatively independent of the ‘sternal stimu that initiate i, This characterises certainly not typical ‘Gt human volitional behavior, although it superficially resembles certain ‘ompulaive behaviors in humans Tvon7 named thie type of behavior —wuhich seems to be nether asin pe reflex nor 2 volional behavior a fixed action patern (FAP). Al though the behavior isnot “xed” in ata sense, this term emphasizes that tis more or less invariable (auggesting a strong genetic component. Table 52-1 compares some ofthe characteristics of reflos, FAPS, and voltional behaviors. Thus FAPs, while being invariable like reflexes, more ‘sey resemble volitional behaviors because of thei steering mechanism However, FAPs diller frm voitona behavior becase they continue after ‘he elcting stimuls is no longer present ‘Any slimulus that ects & FAP—-such a the egg egg-roling behavior or the red-colored “belly” in the attack behavior of the stickleback fish (Figure 52-3)—is called a sign stimulus. For example, a goose will retieve «Wooden egg inthe same way’ as a real epg, so the Wooden egg is also by ‘sfiition a sig stimulus. Yer only the real egg isthe “natura” sign stimu lus, thats, the one that evolved with the behavior. The special ferm re- leaser was originally applied only to the latter category of sign stil. Curiously, “artical” sign stimili, suchas wooden eggs, are sometimes, wore povwerful than releasers. Certain bitds (Figure 52-4, when given & ‘oie, wll hoor a large wooden egg in preference to their own nrmal- Sle eg, In this case the stimulus hasbeen called a super sign stimulus. oe 7A Voto! Bh | Seaton aad saben condoned ‘Animal Beasion 1287 FIGURE 52-2 Egg-oling behav inthe Buropeen graying gone i termed 2 fed econ pate FAP) Qo=— <> G ae particular fate that tigger FPA male sklebsck ah (all ot aac vette model St another male seb it Sacra red Bally), but vl at fick another mde, however ne sie at ae go yh. stimulus, rather ha secogaion (hat ulgges the aggresive behave1238 Chapter 52 FIGURE 52-4 Sometimes bids Sich as this oytreatcer choose Sn sical egg to Incase rather ian thes ow if heart eB (Blneger tims of ti pe now as super sgn sti FIGURE 2-5 The sma or Feeding behavior nt lays pe Seep av evidence by the Seow ware that foding Joung cowed, The feral cw Bin ses no bell nest, but father deposi her gg fo nests of ‘he species, In one well-known ethological model, sign stimu are detected inthe central nervous system by special ensory mechanisms called innate {easing mechanisms CRMs) Each IRMis specific to particular sign sim MRAM us oval-shaped eggs, whether real or aval, are detected by 9 Specie IRM fortis shape The IRM then sts nto motion the FAP nth hse, egg rolling. ‘ cehRr wellknown example of how releasers and FAPS intact i feeding behavior in gulls. A red spot onthe il of certain guls i elo, for begging behavior in nestlings. When the parent gull approaches he fest eth or without food, the IRM inthe young gl tees (oy means ‘Moual stil) the spot onthe parent’ bill and sets into mation a FA. Teihaaly, the begging response. Even a red spot pated on apis ‘EeSboard cls ths begging response. Likewise, the open mouth of ‘ity the pea ofthe re Shiv) ocd ele the four bast ‘eet, ter ate he peak t the lt fn poorer Fiat lo the ight oF st wil gti it slects leathery-hasked and hard-husked fits 25a aye would be worth taveing farther ft meant spending esting Fking, bu of course that would depend on how mack father. ‘Wat do we need to know inorder to slet the best forging skate Cbvioualy we nee fo kno how long If takes fo gel rom one fait ee {he nou Sr ths depends on the deity of the frat es I the dens {ha cess fow tat takes an hour to get rom one lt eet the ‘ont then common sense would tell ws not pass up a harsked fi ‘Retiecnuse takes 10 minutes to husk. On the other hands if te fa IES so dens that i takes only 2 mints to get othe mex te, hen ‘EcTpoutdceainy pass up al the hard-husked forms and petheps wold Tren conser pestng up both the hard-hasked and laherteshed {Doms After the next sol-haske frat s only 6 minutes away, and x takes almost that long just to hask a Rare frit “is teationship between travel ime, handling ime, and net fod in take can be expressed graphically (gure 52-19. This graph, based on {Sgt rather than thre food tem, ndestes eight possible feding ste SRL Resuming that the clonic vale expressed as Ef energy) ofall FER MEoal the best tem would be the ome tha requires the les Nandling doe, the worst would be the item that requires the most hay Ming te, and all others would be ranked accordingly. “Thc muribers onthe horizontal ais do nat fer to the specif tems, bt rather to feeding tees The suber "refers tothe state which the animal slots only the best tem, the one requiring the shortest har ing time (per uni of food and hence the highest rato of energy fo fandting tine (), Yetby Beng so selective, the arma has lage ve Tir (since it has chosen to pass up the ler seven items In the itedingsvategy indicated by 2” the animal selects the best item pus the ‘econ best em, thereby decreasing traveling ne but increasing hat Siig time ond conceuerly decreasing). As te animal Incudes more Sind inne unfavorable food items in te det, travel ime decreases whe Handling time increases. Ths, the animal spends the lest amount of ine veing talc all eight ood stems, ut by adding more untvortie ems tris dct to redace fave! tm, Ht st also spend more tine han dling the tems Te gon hata, where good fod tems ae abundant and the ais doce Soave to evel far te obain these tems, the opal feeding tt gy would ted toward th let of graph (tate 1, 20, depending on good the habitat) In contrast poor haba, here takes nee {0 ial the best fod ites, the optimal atopy woul eto sles or Sema, even though dong s0requces more handling tine How do these cheorteal considerations app tothe actual behavior of animals! By measuring tvel time, handling time, ard eting time 4 ‘Gtng thew values tte theoretical values calelated for opin for {ig one can test the hypothesis that a pata animal does not BR Spually fhe actual vlues do not tthe theoreti vals, 188 ‘Reantthat all elevant variables have not ben accounted for. The mos} {Ray chose to forage seryinecenly (i terms of consuming 8 cae Tuner of lores each day) by selecting oly hare husked frit: Ot Tay do this because it prefer the tate of har-husked ris, oF Bt they contain a cate tent hts absent te ter fmt TH ‘ther hypotheses concerning foraging behavior may’ come o IF ihe monkey deed wes observed foraging optimally, acorn predicted ylucs it may have learned to do sp trough tal and {operant contoning), tats, by randomly tying all tee tae 2 {Cfedng te one that essocsted ih the fewest hunger pain at = othe doy!— a 1 ee MIGRATION ‘nim migration includes two separate but elated phenomena that de- pet om separate physiological mechanisms: (2) the mulus to migrate; td @) orientation and navigation, In many birds, environmental conditions such as temperature, rainfall, length of day, and food availability are so dasly Linked in time to migre- Son that ito tenting to regard these factors asthe “enuses” of migration. Its now thought thatthe stimalus to migrate in most birds is just one vent in a coordinated sequence of events that includes reproduction, olin, and accumulation of fat peor to migration (Figure 52-20) These ‘ents are seen as responses to an innate crcannval hythn (an endoge nous rhythm of approximately one year in duration). The cicanntal Pythm i Based om "iological clock” that has about @ one-year cycle Thisclockcan be “reset duringthe yearby external factors, somewhat ike 2 conventional dock that can be reset throughout the day. The environ- mail stim that reset the cack tthe beginning ofthe migratory season ‘ould be a change in photoperiod, a drop in temperature, or some ather ‘ue. The same envitonmenta stimuli could reset the clock atthe beginning OF the reproductive season. [Navigetion is the abiityof an animal in enfamillar suzoundlings to find is goal without relying on landmarks with which itis famiar. Many inver- ‘bates, including mollusks and arthropods, as well as representatives rote vstually every vertebrate class have exhibited this ability. thas been emeonstrated experimentally tha birds can orient by responding to env- ronmental cues rach asthe position of the sun and stars an the eath’s ogni field. When migzatory birds are placed in drelae cages equipped ith electonically monitored perches they more frequently iy to perches ‘the south side ofthe cage during autumn, One might speculate that they ‘we ryng to escape in the direction they normaly take during migration at Animal Behavior 251 FIGURE 52-20 Seacoal change in the physiology ad bahar of the ‘ehle cow aparow: Thi oo inated sequence of event obese ‘ra oneyear ogi dock”1252 Chapter s2 Teinted 180 degrees, the birds orient toward the north, indicating thee Tome way they use the configuration ofthe stars as directional cues Se ae tage eee ee a a Sy eexeoee ee ae ee a ee an apyree SOCIAL BEHAVIOR Social behavior involves adaptive interactions among animal, espe mong members of the same species. In asessing the extent of soi nave characters ofa given species, ethologsts consider the num nimals that form a group, the portion ofthe lifecycle thatthe grup fentan together, the amount of time and encrgy devoted to socal fn Ton the structure of the group, communication among group ment, formation offmiles, an the nature of interactions. Many ofthe behav ready discussed inthis chopter, including urine marking in dos, a aspect of socal ehavior. Communication “The ability to comumusicat is an essential ingredient of soil behavior, or ‘only by exchanging mutually ecognizable sigals can one animal na the behavior of another (Figure 82-21). Communication occurs whe an finimal performs an act that changes the behavior of another ofan ‘Communication may facitate Finding fod, asin the elaborate dacs the voce It inay hold group together, waa a group of danger, ae Social status, solicitor indiate willingness to provide care, Kenly mm bets ofthe same species, oF indiate sexual maturity “Modes of Communication ‘Animal communication difers significantly from most human cama fionin that itis not symbole. As you ren, information is conveyed War mind by words; yet the words themselves are not the information Tvs and for. The slaionship between the word “eat” and the arin te! isa learned one; person who could ead only Japanese wouldnt re fiz it This snot to say that signals in some sense are not empl! Shims. In a way, ll eleasers are signals. However, relesers ae hecessavily Ieamed, whetens tre symbols ae ‘Although in humans some body communication js culturally de mined lame behaviog, «large part of (such as sing is tray wit fal and appears tobe physiologically determined. The pupil of he wma {ye dates in certain emotional situations, suchas sexual interest or tment. Without realizing it, people respond to such subtle cues. 18 ST sent, 2 photograph ofa voman’s face withthe pupils retouch! pear greatly diated was far more attactive to male subject tha em" tehich they vere shown a8 pinpoints ‘Signals ee often transmitted involuntary as an acompinet physilogial state of the organism, Information about am ani! st onal or mena state may be tansmited even if no other member Species are neat. For example, a bird automaticaly gives 2” ‘hen it sights a predator. Certainly there aze mes when hum Vwould rather not communieate ther true feeings—yet there ™ ae th pi i eth FIGURE 52-21 A mle hyd fog (of Conta Rc ealing tole 2 mate y Sances in which we do not eeally have any choiAnimal Retasior 1253 And Who has not blushed ata tine when he would have given almost ‘ying not to have done 202 "po animals ever employ symbols, or ae anal signal tly restricted wy the equivalent of gasp of alarm? The materi controversial. Many “hologit think that even dogs do ns espond to spoken language swe Sh bat deduce the behavior we command from an sste reading of faiman facial expresion, va intonation, and body atatade. On the ther hand, chimpancwcs have been taught fo spenk a very few words treuingful, and toa ited extent can ds ign language appropriately. Whether apes employ snot nguge in nature ie unbely, although the potential seems to eta. the singing of bis isan obvious example of story communication, serving to announce the presence ofa tertoral mae. Some animals com — fhuncteby scent rather than sound. Anslopes rab the secretions offical FIGURE 52-22 Pronghorn snore ns on copii ebjets nee vent Figure 2-29, Dogs mask RAE ny epg et Terotory by frequent urination, Certain fis, the gymnoids, use electric {05 are ules for navigation and commancatin,inhading erstorlUeat, a Ieshionsimuar to bid vocalization. As £0. Wilson has sai, “The sb, in tect, sng seta song.” Who would have goesed #? Display are visual signs that include movements, postures and faci cnpresions. The comparative study of ada expresions reveals how they {theo to help an ncvial incense, decens, or manana certain tistance fom members ofthe same specs. For ample, the varity of teal expressions observed in member ofthe do fanny (Canidae) ts tutto be unexpectedly large (Table 52-2). Also, elated species fend to ve cnia aca expressions Table 52-2 shows at foxes, which gen Cal have mote felinclke plays than other cand, ifr fom coyote, sles, an dogs nckng the agonistic Bring of teth (vera contac ono pe) eso "A gretr ange of faa expressions is possible when constiuens of displays ae superimposed. Te successive sft in intnsty of fear Sotsion and aggression inthe coyote ane indicated in Figure 52-23, where the positon ofthe ears in 7 is cmbined with the configuration of thesmouth in 3 to produce 9. Th 52-2 COMPONENTS OF FACIAL EXPRESSIONS RELATED TO BEHAVIOR IN CANIDS emer Sete “omponet of Express Aut with TAGeWS rAgewo Md on on nek ete oe errer emia Scam TT ee ee eo eee aan Eee 2 a et tad Rowton Plitit Serer oe a eo sores pe toes Sees mreatrne pe peer Se Bee ier ee See Some aeeeeeees 1254 Chapter 2 a; FIGURE 52-28 shits in intensity {os fesrsebousion (0, 4 7) and Sugeason 23) dog. Face ‘cons ay be emt’ Stpeumposed ain the cae f 9, ‘vlc ls combination of 3 and 7. 3 MA oe a & > atime me Seri Seti nt expen tae ng el res i a ee ee aly sy an oy waarmee eres Sten | creed ace le TE era Sep ee Oe ne ea eager Se ee ee Soo eee ee acm t | ey Dominance Hierarchies ‘quale among themehves by chasing and pecking each other untl Mibtich a nk onder, or dminance hierarchy, among theme {navidad atthe fop of this dominance hierarchy dominates al the | ‘duals In some species the daminance hierarchy i sad tobe lines wt | {he dominant individual being represented by a letter suchas At 219, | oesachy, A dominates over the next highest aking individu B 2° ls over all thers in the group; B dominates over Cand all others ben | and so forth, However, rank order in some groups isnot ines ‘dominating over B, Bover C, and Cover A. In larger groups he | ‘more complex hierarchies | Many years ago it was observed that chickens living together i8 2 5° indPee Confit that reinforce rank order may occu when individuals compete {or food in the tough, Dominant individuals win most but not necessarily of their encounters with subordinate individuals. tn many vertebrates fighting rarely occurs once rank has been established because individuals {quick Tear ohich members of the group are above them in rank (Figure ‘3-29. When the dominance hierarchy ofa group of animals i studied in depth, questions such asthe fllowing come to mind, 1. Which physical characteristics influence an individual's rank in the bicrarchy? 2, Does an individ’ past history influence its rank in the hierarchy? 3. Does the rank of an individuals parents influence that individva’s position in the hierarchy? 4, Kre contests between individuals more frequent at certain levels in the hierarchy? 5. Isthere a correlation between the rank of an individual an its repro- ductive success? In general, dominance hierarchies have been interpreted as adaptive be- fuse they make it unnecessary to expend energy for continually compet ing with group members for food resources, Teritoriality Virtually all animals, and even some plant, maintain a minimum personal Aislance from their neighbors, as one can observe in the even spacing mong the members ofa flock of birds resting on a telephone line: Most {nimals havea geographical area that they seldom or never lave, Such an tren'scalled home range (Figure $2-2). Sine the animal has the oppor tunity fo Become familar with everything in that range it hasan advan- ‘age over both its predators and ts prey is negotating cover and tinaing food. Some, but not al, animals defend a portion of the home range gainst other individuals ofthe same species and even against individuals tfother species, Such a defended areas calle a teriory. The tendency to Akefend sch 8 teritory Is known as trsitoriality. Terntovality i easily studied in beds. Typically, the male chooses a lentory at the beginning of the Breeding season. This behavior results ‘fon high concentrations of sx hormones inthe blood. The males of adja ‘tnt tentores fight unl tertorial boundaries become fixed. Generally, the dominance ofa cock varies directly with his nearness to the center of his tertory. Thus, close to “home” he sion, When invading some other ‘nds ternitory, he is Hkely tobe a lamb. The interplay of dominance values among territorial cocks eventually produces a neutral line at which nether indominant. That line i the teritoril boundary. Bird songs announce the lxistence ofa territory and often serve asa substitute for violence, Further "mote, they announce to eligible females Wat a propertid male resides in ‘he tersitry. Typically, male bids take up a conspicuous station, sing, and Sometimes display siking pattems of coloration to their neighbors and Tras (Figure 32-26) “Temntorality among animals may be adaptive in that it tend to reduce conflet,contrl popslation growth, and ensure the most efficent use of frvitonmental resources By encouraging dispersion and spacing organ isms more or las evenly throughout a habitat. Usually tevtorial behavior 'Srolted to the specie Mf-style ofthe organism that displays it, and to Ihatever aspect ofits ecology i most eit! to its reproductive success, instance, sea birds may range over hundreds of square miles of open rater but exhibit testoril behavior thats restricted nesting sites on 3 "oe or island, their resource tha sin the shorest supply and for which ‘competition is keenest. Animal Beasfon 1255 ‘many signals to convey messages Felt focal omifance te the ceo aba, ne sgl snes ‘Stok toward male osigaly ‘ness to mate, Subordingte tales eth sme pete 10 a5 ‘ine dominant ue that they do bot intend to een te higher Taking sna Ha a suber ‘ate mle present fo 9 nore domi: ‘am resaures the subordinate with osc pt om the Back FAURE 38 A cor we a trary secaded aes in hich 2 teens on esas ome ange. Among Se most ter font of er! rea ees he sory eal lured hee whch ‘oa aac ay smal ean = Tian diver tat comes too dose toite sel1256 Chapter 82 FIGURE 52-26 Courtship displays. (Ate weed a en {fe Galapagn inland (h) Cour Sis by meer es Speco exch specs. Ths Fiala sequence ofthe motion the large gt vei hace Buc ofthe pees Ua ee. FIGURE 52-27 Male and female Fumping spiders, Pes cus, Ine ership har tat pre {Eucs mating ema procs Sorte dance tht ss the ‘ai’s tural aggresion toward ‘allowing hat fo get ose ‘hough to neeminate her Sexual Behavior and Reproduction “The minimum socal contact and fr some species of animals (for ample, ‘many species of spiders) the ony social contact, isthe sex act. Fertilization land pethaps the fearing of young are some animals’ only forms of scl ‘behavior (igure 52-27). Let us consider the sex act a abasic example of Solel Behavior, forthe elements to which it can be reduced are also the least common denominators of most soil behavior. Tike other socal eelatonships, the sex acts adaptive in that it promotes the welfare of the species Ie requires copeaton, he tepory suppres gprs bohvor, and ete of communication. Among some jumping ‘Spiders, fr example, mating s preceded by 8 rital courtship on the part ot the tale, the efiect of which ito produce temporary paralysis in the f inate. While she i thas enthralled, the male inseminates her. Shou she ‘ecover before he makes his escape, he comes the main course at his ‘wedding feast, Whether he appreciates the opportunity or no, he i ts thle to ke the ultimate in material contsbutions tothe eg the female Will presenily produce, She would otherwise have to bear the metabo burden of their production all by herset ‘Since an individual that reproduces perpetuates its genes it is nots prising that natural selection has favored mechanisms including bs that promote sucessful reproduction. To fertilize as many females = Pr sible, the males often compete intensely with ane another. Sexal compet ton emong males ofthe same species often has contributed tothe cv tion of large tale size, brillant breeding colors, omaments, anes a4 folk features that give a male an advantage in establishing dominan™ fmong his peers and attracting females. Since the female usually chooses the mate selection has favored thst male characteristics that make a male most attractive. Selection his 2 favored those female atibates that enable her to determine that he is worthy of her investment, Success of male in dominance encom tvth other males indicates his tness othe female. Athougl the fh ‘ame species accepts the frst male that attempt to court hes Species the female tests the males by provoking encounters, Fert Hons and chimpanzees in ests have enlarged, bellatl closrllings that atract all males and incite competition among them (Figure 2-28), The victorious male courts the female, An important function of court ships to ensue that the male is member of the same species, butt also provides the female further opportunity to evaluate the quality af the male Courtship may also be necessary asa signal to tigger est building oF ‘ovulation. Courtship ntuals may be long and complex. The fist display of the male releases a counter behavior of conspectic female. This, i tar, rslenses additonal male behavior, and so on unl the pair are ready for {npulation. Certain male spiders make an offering of fod to the female ‘Eurng courtship. This inhibits any aggressive tendencies thatthe female may have on Being approached and also provides the female with some of the food needed for egg production. ‘Sexual selection has also led to strategies whereby a succesful male protects an inseminated female from copulation with other males. After {npulation a male damselfly continues to grasp and fly with the female Unt she has deposited her eggs. A succesful drone honeybee discharges rch of his genital apparatus into the virgin queen's genital passage, Thereby blocking them against insemination by another male Pair Bonds | pair bond isa stable relationship between animals of the opposite sex that ensures cooperative behavior in mating and the rearing ofthe Young (Figure 52-29) In some species a newly arsved females ntl rated as 2 rval male. Then, through the use of instinctive appeasement postures and gestures by both male and female, the inital hostity is cissipated ana ‘mating takes place. Such sexual appeasement behavior may be very clabo- tale ned iors vi: to mating dane fn som ide “The releaser mechanisms involved in the establishment and mainte- tance ofthe pair bond are often remarkably detailed, A male ficker pos- Sesies a black, mustache-hike marking under the beak. This is lacking in the female If “happily marred” female flicker is capture and such a ms- "athe i painted on her, her mate wil vigorously attack her as ifshe were a ‘val male. He will accept her again if itis removed. Such cues enable Courtship rials to function aa behavioral genetic isolating mechanisms mong species Care of the Young Cre ofthe young is an ational component af successful reproduction in many species, and i oo, requires a parental investment (Figure 52-30} The benefit of parental eae is he increted likelihood ofthe survival ofthe cifepring, bat the cost sa reduction in the numberof offspring that can be ‘rodtuced. Because ofthe time spent carrying the developing embryo, the female has more to lose than the male f the Young do not develop. Thus, females are more likely to brood eggs and young than mes, and asvally the females invest more in parental car, Tnvesting time and effort in cate of the young is usually less advanta- ‘tou toa male, for time spent in parenting is ime lst in inseminating ster females. Even worse, It may not be certain who fathered the off Spring. Raising some other male's offepring is definite genetic disadvan- lige. Tn some situations, however, it may be to the male's advantage to help ear his own young, or those ofa genetic relative. Receptive females maybe saree. And gathering sulicient food may requice mote effort than one paent can provide. In some habitats the young may need! protection “inst predators. Animal Behavior 1257 FIGURE $2-28 This female baboon Etgals pedi sexual readiness she bight red postenor FIGURE 52-29 A prof nest slstrosses In uy specs pale Sons ae maintain (rover dapaye of Ey grming1258, FIGURE 52-30 Examples of parental svestren (Eh Cougars and bac beer are normally orl enemy (arsed ovoid cach ether, Ths contontation ya ‘Biated when the cougar ited i the ares whe 9 [Emte bear ws sing her cubs.) A chistap pe tern eprmng an re you, Such on es. Sten of tikes energy onthe pat of he patent cs poten ips yd lp ne nso ofthe pent gees no sutceeding bee ‘Re A baby baboon aides ont mothers be dye ig en ivancy and comes to ines ome fe Play Py is an important aspect ofthe development of behavior in many spe ‘ies especially young tnannas, It serves asa means of practicing ait piltem of bchavior and perfecting means of escape, prey killing, an even FEwual conduct In true play the behavior may not be actully consun: ‘mated, Thus a kifen pounces upon «dead leaf but ofcourse doesnot {even though the kitten adainisters atypia carivore neck bite. When playing with liermate, the same kitten may practice the disemboweing Evoke with its hind claws (Figure 52-31), but the littermate fs not ner tionally injured inthe process. Sociobiology Sacicbology isthe school of ethology that focuses the evaitono Socal behator tough natural selection, Like many biologi fhe ps {fading Darin, Edward. Wilson and ther scitiolgis em sir te sina! rots of human behavior, but they ave attempted 1 fun er dlcpine wih population gence, wih paral emp o* the eet a on sletion (Chapter 19) on puters of inhestane ar the acnbslgat, the orgaiam ands adaptations inci behavion=-ave ways genes have of making more copies of thems Th cells ane aes of he body support the funetions of the reproduc ‘sem: The reproductive sts jabs the tanamision of genet mon cing gon : . here ae ungue peas in stmptng to reconstruc the eve beaior, since Esko arly eve an expt foal ecard. i atinally, by applying human socal terms to behavior in animals that may be only superficially similar, we create the perhaps entirely false impression ‘hat itis the same behavir. Ie isan easy step ftom that f the assumption that the causes and utility ofthese behaviors are the same as those of torresponding human behavior. Consider, for instance, the question of ‘whether humans ae teritorial. We do tend to preserve space between us 2s individuals, to defend our homes, and as groups to defend larger, poi ‘alareas. However, do these bchaviors have the same genelicand adaptive Value in humans asin animals? And is human tertorialty homologous With that of other animals, oF is it mercy analogous? Also, problems of objectivity can est. Any sumptions we may have shout our own territoriality can cause us fo look athe behavior of animals ssa miror of our own, Among closely related species of primates, socal ‘organization and the degre of tertriality and aggressive behavior vary widely. Which of these species should we choose as models for studying hhuman behavior? ‘Most of the controversy that has been tiggered by sociobiology seems ‘elite to its possible ethical implications. Sociobiology is often taken 25 ‘denying that Ruman behavior is leble enough to permit substantial provements in the quality of our social lives. Yet socobologists do not ‘sagree with their eric that human behavior is flexble. The debate therefore seems to rest on the dee to vehich human behavior is genetic andthe exon to which i con be modified ‘As sociobologists acknowledge, people through culture possess the ability to change their way ofife far more profoundly afew yeas than 3 hive bees ora troop of baboons could accomplish in hundreds of genera oes of genetic evolution, This ability Is indeed genetically determine, nd that is a very great gift. How we we tan what we accomplish with i ‘sot a gift but a responsibilty upon which our own well-being and the wellbeing of other species depend, SUMMARY ‘Animal Behavior 1259 FIGURE 52-31 Young ons playin Inouthern Ai: Py behtioe thats not consummated and often ‘aha that wl be Ged in est in ater Me, posi in ne Ing. Sting for rary, or com Paling fr tates. {Animal behsvios, or the movements of animals n= I. The questions that nimalbehaviests sk nally ll onset stnul, may be studied frm the points o€ into ene of to eaegunes, er of several subicpines ‘A. Questonsconceed with longterm evhionary [A Chass ethology focuses primary on the char ‘ues of behavior, elle timate factors, steric, adapeve vale, and evlutonary i '. Questons concerned with immediate conditions {ory of inate, species! Behavior 3B. Bahavioe genetic i concer with how behav- of mechanism, called proximate factors, Central to much of cles ethology the Give lor rite ae inert, ‘con patem (FAD, an innate, stereotyped sponse ©. Comparative peyehology typically deals with faa simul (a sig simul, of let tates learned Behavior and Row It can be priced fase component and that continues ler the stl ‘nderspeciid conditions, fas cea . Behavioral eccogystudes behavior incelaton to A Sign simul ate detected in the central nervous ecological variate. spit by special sensory mechanisms known at , Neurobiology seks to explain behavior mech Siete resting mechan, cath of wich sms in anata ane phyogs spelctoa partculr