Author Institution: Department of Biology, Wilmington College ; Department of Biology, Earlham CollegeTerritorial aggressive behavior was studied in male brook sticklebacks collected in Saskatoon, Saskatchewan; Oshkosh, Wisconsin; Ft. Atkinson, Wisconsin, and Urbana, Ohio. In the 20 h of observation 1,167 individual encounters with 3,305 separate aggressive displays were observed. Aggressive behavior was observed to be complex with at least 12 distinct aggressive display postures observed

Burks, D. J.

Scantland, D. A.

Sinclair, R. S.

Woolpy, J. H.

The Ohio Journal of Science

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Copyright © 1985 Ohio Acad. Sci. OO3O-O95O/85/OOO1-OO23 $2.00/0GEOGRAPHIC VARIATION IN AGONISTIC RESPONSESOF TERRITORIAL MALE BROOK STICKLEBACKS,CULAEA INCONSTANS1DOUGLAS J. BURKS, Department of Biology, Wilmington College, Wilmington, OH 45177D. ALLEN SCANTLAND, R. SCOT SINCLAIR, and JEROME H. WOOLPY, Department of Biology,Earlham College, Richmond, IN 47374ABSTRACT. Territorial aggressive behavior was studied in male brook sticklebackscollected in Saskatoon, Saskatchewan; Oshkosh, Wisconsin; Ft. Atkinson, Wisconsin,and Urbana, Ohio. In the 20 h of observation 1,167 individual encounters with3,305 separate aggressive displays were observed. Aggressive behavior was observed tobe complex with at least 12 distinct aggressive display postures observed.Fish from Saskatoon, Oshkosh, and Ft. Atkinson demonstrated several similarities intheir territorial behavior. Charging was the most frequently observed display in all threepopulations. Also, biting, sigmoid and broadside displays were within the top four mostfrequent displays in all three populations offish. The relative frequency of all aggressivepostures were similar. Analysis of following event display frequencies (the frequency thatone display follows another) also showed striking similarity in the behavior of territorialmales from Wisconsin and Saskatchewan.The Urbana population varied significantly from the other populations. The Urbanafish showed fewer postures, had shorter encounters, and did not display any attackpostures (agonistic displays which typically lead to an attack or bite).OHIO J. SCI. 85 (1): 23-29, 1985INTRODUCTIONThe stickleback family, Gasterosteidae,has been the subject of rather extensivebehavioral studies. Aggressive behaviorhas been studied in the threespine stick-leback, Gasterosteus aculeatus (van denAssem 1967, Huntingford 1981), in andbetween the fourspine stickleback, Apeltesquadracus, and the threespine stickleback(Rowland 1983) in Pungitius pungitius, theninespine stickleback (Hoogland et al.1957, and Huntingford 1977), and inthe brook stickleback, Culaea inconstans(McKenzie 1969a + b, Reisman andCade 1967.)The brook stickleback is the only com-pletely freshwater North American stick-leback. It has a rather broad range inNorth America. It is found in ponds, lakesand streams from British Columbia to New'Manuscript received 18 May 1984 and in revisedform 21 December 1984 (#84-30).23Brunswick and from Nebraska to NewYork (Nelson 1969). Fish are observed tovary in several important morphologicalcharacteristics within this range. Signifi-cant differences in dorsal and pelvic spinelength and in body depth have been ob-served (Nelson 1969). Spin length is gen-erally greatest in the Ohio/Wisconsinregion and is observed to decrease as onemoves northwest within the range. Alsothe pelvic skeleton is largest in proportionto the body size in fish observed in theWisconsin to New York populations andleast developed in the northwestern popu-lations (Nelson 1969).The observation of clinal variation inmorphological characteristics of the brookstickleback led us to investigate the behav-ioral responses of sticklebacks in relation togeographic distribution. In this study wereport results on the agonistic behavior ofterritorial male fish from populations inOhio to the Northwest area of Saskatoon,Saskatchewan, Canada. Males showed a24 BURKS ET AL. Vol. 85very complex pattern of territorial aggres-sive behavior. The Urbana, Ohio fish variedmost morphologically and in their behav-ior from the other populations studied.The fish from Saskatoon, Saskatchewan;Oshkosh, Wisconsin, and Ft. Atkinson,Wisconsin, showed surprising similaritiesin behavior considering the complexityobserved in aggressive behavior.METHODS AND MATERIALSFish were collected or purchased from Saskatoon,Saskatchewan; Ft. Atkinson, Wisconsin; Oshkosh,Wisconsin, and Urbana, Ohio (fig. 1). Prior to andthroughout the duration of the study, environmentalconditions were maintained at the optimum level foreliciting territorial reproductive behavior (WinnI960, Reisman 1961, and Smith 1970). Since thesexes are difficult to distinguish until the re-productive cycle begins, eight to 12 fish from onearea were placed in a 115-X-40-cm, 120-1 tank.Reproductive males were selected for the study andfour reproductive males were left in the tank. Thetanks had plants and rocks to serve as cover alongwith material for nesting. Males used in this studyshowed dark skin coloration, black bar eyes andnest-building behavior.The fish were observed through 4-X-10-cm slitsin a black plastic screen hung in front of each tank.Fish from each area were observed for a total of five heach. The frequency and order of occurrence of thedifferent display patterns were recorded. Ob-servations consisted of 30-min periods duringwhich the observer watched two territorial fish in0 250 500 750FIGURE 1. Range and collection sites of malesticklebacks (Trautman 1957).the same tank for 15 min each, while another ob-server recorded these data.DESCRIPTION OF AGONISTICBEHAVIORAL DISPLAYSFRONTAL APPROACH (fa) — The dis-playing fish moves towards the other terri-torial male with its head pointing towardsthe intruder.BROADSIDE (bs) — The territorial fish ori-ents itself at a right angle to the intruder.LATERAL DISPLAY (la) — The territorialmale orients itself in a side-by-side mannerwith the other fish typically with its headtowards the other fish's tail.SIGMOID (s) — The displaying fish con-torts its body into the form of an "S" itshead directed at the other fish.SIGMOID ATKINSON (sa) — A low in-tensity "S" in which the territorial male'sbody is bent into only half an "S."HEAD DOWN (hd) — As a reproductivemale approaches another fish, its headis positioned down at approximately 45°angle.TAILBEATING (tb) — A display in whichthe fish sways or beats its tail side-to-sideat a rapid rate.FOLLOW (fo)— A fish swims slowly inthe direction of a fleeing fish. This appearsto be a low intensity charge.CHARGE (ch) — The displaying fishdarts at another.CIRCULAR FIGHTING (cf) The two fishchase each other in a circle that has a di-ameter of five to 10 cm. The chasing ishead to tail.BITING (bi) — The displaying fish bitesthe other fish.DRAGGING (dg) — After a bite the dis-playing fish holds onto the other and dragshim for some distance.TERMINATION (te) — The fish termi-nates the encounter by fleeing or by nolonger participating in aggressive display.RESULTSTERRITORIAL BEHAVIOR. The total num-ber of aggressive displays range from a lowof 25 displays observed in males collectedfrom Urbana, Ohio, to 1,724 displaysOhioJ. Sci. STICKLEBACK BEHAVIOR 25TABLE 1Number and length of aggressive encounters betweenterritorial male sticklebacks.LocationUrbana, OhioFt. Atkinson,WisconsinOshkosh, WisconsinSaskatoon, CanadaNumberof Displays258376181729Numberof Encounters18297168593TABLE 2Average number of displays per encounter betweentwo territorial males.AreaAverage Number ofDisplays Per EncounterUrbana, OhioFt. Atkinson, WisconsinOshkosh, WisconsinSaskatoon, Canada1.382.823.682.92within the five-h observation period in theSaskatoon, Saskatchewan, males (table 1).The number of displays observed per unittime appears to increase as one moves fromOhio to the northwest part of the range ofthe brook stickleback (table 1). In somepopulations not all of the range of ag-gressive displays were observed.The number of displays in an encounterranged from one up to 15. The averagelength of an encounter ranged from1.38 displays per encounter in the Ur-bana, Ohio, fish to 3.68 displays per en-counter in the Oshkosh, Wisconsin, fish(table 2). There were no statistically sig-nificant differences in average encounterlength between any of the populations.In terms of the frequency of individualdisplay occurrence within a group, it wasobserved that the Saskatoon, Oshkosh, andFt. Atkinson fish showed several simi-larities. The charge was the highest fre-quency display in all three populations.The broadside, sigmoid, and bite were thenext most commonly observed behaviors.Variation was observed in the relative fre-quencies of these behaviors among thethree populations. However, the simi-larities between the three populations arestriking. In the Urbana, Ohio, fish onlythree behaviors were observed. Two of thedisplays (broadside and sigmoid) werein the four most commonly observedbehaviors in the three other populations(table 3). The third display (lateral dis-play, la) was a high frequency display inthe other three populations.Each half-hour observation period con-sisted of observing two different male fish,each for 15 min. Table 4 shows the com-parison of two fish and the frequency atTABLE 3Frequency of individual display postures within a geographic group.Frequency12345678910Urbanabs* (40%)la (36%)s (24%)—Ft. Atkinsonch (49%)bi (18%)s (10%)bs ( 6%)cf ( 5%)la ( 4%)fa ( 4%)hd ( 2%)sa ( 1%)tb ( 1%)Oshkoshch (25%)s (20%)bs (17%)bi (10%)la ( 7%)hd ( 6%)fa ( 5%)tb ( 5%)cf ( 4%)fo ( 2%)Saskatoonch (48%)bs (17%)bi (16%)s ( 6%)fa ( 5%)fo ( 4%)la ( 3%)hd ( 1%)tb ( 1%)cf ( 1%)Nbs — broadside, ch — charging, la = lateral display, s = sigmoid display, bi — biting, cf — circularfighting, fa — frontal approach, hd = head down display, fo = follow, tb = tailbeating, sa = sigmoidatkinson, dg = dragging, te = termination.26 BURKS ET AL. Vol. 85TABLE 4Frequency of individual display occurrence comparingtwo fish from the same collection point.Saskatoon ach*bsbifoslafa38.0%26.6%12.4%10.4%6.0%4.7%1.9%Saskatoon bchbibsfolastbfacf42.0%22.0%19.0%3.0%3.0%3.0%3.0%2.9%2.0%Ft.schhdbsbilaAtkin. a43.0%27.0%15.0%6.0%4.0%4.0%Ft. Atkischhdbstb41.19,17.17.n. b.0%,0%.0%.0%6.0%"bs — broadside, ch = charging, la — lateraldisplay, s = sigmoid display, bi = biting,cf = circular fighting, fa = frontal approach,hd = head down display, fo = follow, tb = tail-beating, sa = sigmoid atkinson, dg = dragging,te = termination.which various displays were observed.These data indicate that the variancebetween two fish from the same collec-tion point was as great or greater than thedifferences observed between fish col-lected from Oshkosh, Ft. Atkinson, andSaskatoon.Another way to analyze sticklebackaggressive behavior is to determine thefrequency with which one aggressive be-havioral posture follows another. Whenthis was done, several interesting obser-vations were made. The Wisconsin andSaskatoon fish showed many similarities,even though their behavioral displays werecomplex. The Urbana fish were strikinglydifferent, again because of the low fre-quency of encounters (fig. 2 and table 5).DISCUSSIONClearly the territorial behavior of malebrook sticklebacks is more complex thanpreviously thought. We observed 12 dis-tinctive aggressive displays. One displayhas not been described previously. We havenamed this display the "follow." In thisdisplay a territorial male slowly follows anintruding male out of his territory. It isTABLE 5Following event frequency; twenty most frequent display pairs.Population:Urbana Ft. Atkinson Oshkosh Saskatoonla*bssbslalate 6(24%)te 6(24%)te 6(24%)s 4(16%)bs 2(8%)s 1(4%)chchchbibisfobscfcfsbibilalabschfasfachbitetechtetetechbibscfbiteschcftechch155(18%)119(14%)109(13%)63(7%)61(7%)42(5%)24(3%)18(2%)16(2%)16(2%)13(1%)13(1%)12(1%)11(1%)11(1%)10(1%)10(1%)9(1%)9(1%)8(1%)c h ^ chch-H> bich —> tebi —> tes •chlahd-bs-bs-b s •s •s •s •s •b s •b i •bs-bs-bi •tessteschtetbscfbslachhdbsbi42(7%)39(6%)36(6%)32(5,%)31(5%)26(4%)23(4%)19(3%)19(3%)19(3%)19(3%)18(3%)18(3%)18(3%)15(2%)12(2%)11(2%)10(2%)10(2%)9(1%)chchchbsbibisfafobschbifalabsslabslas270(16%)265(15%)249(14%)206(12%)162(9%)77(4%)50(3%)42(2%)42(2%)38(2%)25(1%)20(1%)17(1%)14(1%)14(1%)13(1%)12(1%)12(1%)11(1%)11(1%)"bs — broadside, ch = charging, la — lateral display, s = sigmoid display, bi = biting, cf — circularfighting, fa = frontal approach, hd = head down display, fo — follow, tb = tailbeating, sa = sigmoidatkinson, dg = dragging, te = termination.Ohio J. Sci. STICKLEBACK BEHAVIOR 27B100%FIGURE 2. Following event frequency. This figure lists the frequency at which one display was followedby another. A. Oshkosh, B. Urbana, C. Ft. Atkinson, D. Saskatoon, bs = broadside, ch — charging,la = lateral display, s = sigmoid, bi = biting, cf = circular fighting, fa = frontal approach, hd = headdown, fo = follow, tb = tailbeating, sa = sigmoid atkinson, dg = dragging, te = termination.28 BURKS ET AL. Vol. 85easily differentiated from a charge. Boththe charge and follow display were ob-served in the same fish during a 15-minobservation time period. The follow wasobserved in fish from all but the Urbanapopulation. In the Ft. Atkinson fish weobserved another unique behavior whichseemed to be a variation of the sigmoidposture. This appears to be a low intensitysigmoid in which the fish bends its body ina partial "S." Individual Ft. Atkinson fishwere observed to use both postures.The Oshkosh, Ft. Atkinson, and Saska-toon fish showed many similarities in be-havior. Frequency and frequency order inwhich displays were observed were similarin all three populations. The charge wasthe most frequently observed display inthese three populations. The next mostcommonly observed behaviors were thesigmoid, broadside, and bite. With thesethree displays, the order was slightly dif-ferent in each population. Even as one goesdown the rest of the list ranking frequencyof individual displays, one observes manysimilarities in the three populations. Thevariation in behavior in the three popu-lations seems no greater than betweenvarious individuals within the same popu-lation (tables 3 and 4). Encounter lengthwas similar in all three populations. Theaverage encounter length varied from 2.82to 3-68 (table 2). Following event fre-quencies were similar to the extent that ourstudy permits analysis. The greatest vari-ation between populations was the numberof displays observed during the five-h pe-riod. A three-fold difference in the numberof displays observed was found.The behavior of the Urbana fish variedsignificantly from the other three popu-lations studied. The Urbana fish showedthe fewest displays, fewest number of dif-ferent displays, shortest encounters andvery few attack postures (charging, biting,and dragging). The Urbana fish showedthe greatest difference in morphologicalcharacteristics when comparing the fourpopulations. A distinct difference in spinelength in the Urbana fish was observed.The spines were larger. Spine length wassimilar in fish from the other three popu-lations studied. Also, territorial malesfrom Urbana were significantly lighter incoloration than other territorial fish. Wecould not distinguish by color males fromthe other three populations. It is not clearif the difference in coloration of the Urbanaterritorial males accounts for the signifi-cant differences in aggressive behaviorobserved within this population. This pos-sibility needs to be studied in the brookstickleback to see if it is similar to thethreespine stickleback in this regard (Pelk-wijk and Tinbergen 1937). Another possi-ble explanation for the dramatic differencein the behavioral patterns found in theUrbana fish is their geographical isolationfrom other populations (Trautman 1957).Further work is needed to determine if thedifferences in behavior observed in the Ur-bana fish is a reflection of clinal variationin morphology (Nelson 1969) or if it is theresult of their geographic isolation (Traut-man 1957). More populations from Ohio,Michigan, and Wisconsin need to bestudied.ACKNOWLEDGMENTS. We would like to thank S.Fredrik Anliot for his careful reading of this manu-script and for his many helpful suggestions. Wewould like to thank Carole Lee Kenney for carefullytyping this manuscript and Dorothy Gurtzweiler forpreparing the figures for this paper.LITERATURE CITEDHoogland, R., D. Morris and N. Tinbergen1957 The spines of sticklebacks (Gasterosteusand Pygosteus) as means of defense against preda-tors (Perca and Esox). Behaviour 10: 205-236.Huntingford, F. A. 1977 Inter- and intraspecificaggression in male sticklebacks. Copeia 1977:158-159.1981 Further evidence for an associationbetween lateral scute number and aggressivenessin the threespine stickleback (Gasterosteusaculeatus). Copeia 1981: 717-720.McKenzie, J. A. 1969a The courtship behaviorof the male brook stickleback, Culaea inconstans.Can. J. Zool. 47: 1281-1286.1969b A descriptive analysis of the ag-gressive behavior of the male brook stickleback,Culaea inconstans. Can. J. Zool. 47: 1275-1279.Nelson, J. S. 1969 Geographic variation in thebrook stickleback, Culaea inconstans, and notes onOhioJ. Sci. STICKLEBACK BEHAVIOR 29nomenclature and distribution. J. Fish. Res. Bd.Can. 26: 2431-2447.Pelwijk, J .J . and Tinbergen, N. 1937 Einereizbiologische analyse einiger verhaltensweisenvon Gasterosteus aculeatus. L. Z. Tierpsychol. 1:193-200.Reisman, H. M. 1961 The reproductive behav-ior of the five-spined stickleback, Eucalia incon-stant (Kirkland). Amer. Zool. 1: 468.and T. J. Cade 1967 Physiologicaland behavioral aspects of reproduction in thebrook stickleback, Culaea inconstans. Amer. Midi.Natur. 77: 257-295.Rowland, W.J. 1983 Interspecific aggressionin sticklebacks: Gasterosteus aculeatus displacesApeltes quadracus. Copeia 1983: 541-544.Smith, R.J.F. 1970 Effects of food availabilityon aggression and nest building in brook stick-leback {Culaea inconstans). J. Fish Res. Bd. Can.27: 2350-2355.Trautman, M. B. 1957 The fishes of Ohio. OhioState Univ. Press, Columbus. 617-618.van den Assem, J. 1967 Territory in the three-spined stickleback, Gasterosteus aculeatus. Behav-iour Supplement 16: 1-164.Winn, H. E. I960 Biology of the brook stick-leback, Eucalia inconstans (Kirkland). Amer.Midi. Natur. 63: 424-438.

Geographic Variation in Agonistic Responses of Territorial Male Brook Sticklebacks, Culae Inconstans

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Geographic Variation in Agonistic Responses of Territorial Male Brook Sticklebacks, Culae Inconstans

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