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P0508-P0518.Pdf INTERSPECIFIC AGGRESSIVE BEHAVIOR OF THE POLYANDROUS NORTHERN JACANA (JACANA $PINOSA) MARTIN L. STEPHENS Allee Laboratoryof AnimalBehavior, The Universityof Chicago,940 East57th Street, Chicago,Illinois 60637 USA ABSTRACT.--Adultsof the polyandrous Northern Jacana(Jacana spinosa) defend their off- spring against avian territorial intruders that are potential offspring predators. I investigated this defensive behavior in CostaRica during 1980 and 1981. Jacanasattacked 15 species;most attackswere against Purple Gallinules (Porphyrulamartinica), which are known predators of jacana eggs and offspring. Three factors influenced the probability of attacks:(1) spatial proximity of intruders to jacanaoffspring, (2) speciesidentity of intruders,and (3) stageof breeding (jacanaswere most responsive to intruders when young offspring were present). Attacks prevented intruders from reaching jacanaoffspring, and limited evidence suggests that attacksalso reduced the density of intruders near jacana nests. The female's role in offspring defense was substantial. Females participated in the defense of eggs and small young at levels comparableto thoseof males.This participationincluded: (1) joining ongoing attackswhen their mates solicited aid, (2) continuing attackswhile their mates led offspring away from intruders, and (3) launching attacks in the absence of their mates, especially during incubation. Received8 November1982; resubmitted 19 September•1983, accepted 12 January 1984. NORTHERNJacanas (Jacana spinosa) have one RodriguezNational Wildlife Refuge,formerly known of the rarest breeding systemsknown among as Hacienda Palo Verde, in Guanacaste Province, Costa birds: resource-defensepolyandry (Jenni 1974, Rica.The study site is part of a marshnear the Temp- Emlen and Oring 1977, Graul et al. 1977). Fe- isque River. The marsh is an expanseof emergent vegetationinterrupted by open water and floating male Northern Jacanas defend territories that vegetation. Scattered"Palo Verde" trees (Parkinsonia include the smaller territories of their one to sp.) give the marsh a savannah-like appearance. four mates.Females are nearly 70%heavier than Hundreds of jacanasarrive for breeding at the marsh males and do virtually no incubating, brood- when it is inundated during the rainy season(May- ing, or escortingof the precocialyoung (Jenni November; Slud 1980). The study site was covered and Collier 1972, Jenni and Betts 1978). Fe- by floating vegetation(primarily Nymphaea,Eichornia, malesdo aid malesin defending offspringfrom and Neptunia)and borderedby emergents(primarily predators,however: males and females both at- Typha, Panicurn,Paspalum, and Eleocharis).Small tack birds of other species that are potential patchesof emergentsdotted the zone of floating vegetation. predatorsof jacanaeggs and chicks. Jenni and I conducted observations from towers 3-6 m tall Betts(1978) describethe behaviorsthat jacanas that were within the territories of focal animals. I employ in this interspecificaggression, docu- draped a blind over the tower before beginning sys- ment the time that malesand femalesspend in tematic observations.My approach to and ascent of attacking offspringpredators, and comment on the tower sometimes caused nonfocal birds to flee, other aspectsof this behavior. In this paper, I including speciesthat jacanasattack. In order to re- confirm and extend the observations of Jenni duce any effectsthis might have had on the data, I and Bettsand place emphasison: (1) the prox- waited in the blind at least 5 min, and often at least imate causes of interspecific attacks, (2) the 20 min, before beginning to sample. consequencesof such attacks,and (3) the role During 1981, observationperiods were 2 h long. that femalesplay in this form of parental care. During each period I simultaneouslyobserved one female and one of her mates, regardlessof the num- ber of mates the female had. I observed the same METHODS mate during the entire period over which a particu- lar female was observed,except when events such as I studied Northern Jacanasfrom June to November clutch loss made it profitable for me to initiate ob- 1980 and May to December 1981 in the Dr. Rafael servations of a different mate and discontinue obser- 508 The Auk 101: 508-518. July 1984 July1984] JacanaInterspecific Aggression 509 vations of the first. The focal male and female are causeof the gradual transition from downy to juve- referred to as the "focal pair," even though some nal plumage. focal females were polyandrous. Nonparametricstatistical tests employed below are During 1980,observation periods were 80 min long. describedby Siegel (1956). When meansare strongly During each sample ! simultaneouslyobserved one affected by outliers, medians are reported. An as- female and one of her mates for 60 min; then, after sumption of many of the statisticaltests that follow a 5-min pause,! observedthe same female but a dif- is that attacks are temporally independent of one ferent mate for the remaining 15 min. Males of mo- another. ! determined that this was indeed the case nogamous females were observed for the entire pe- by plotting interattackintervals as a log survivorship riod. The same male was observedduring all 60-min function for each speciesthat was attacked and by segmentsof the period spent observing a particular comparing this distribution to the one that would be female, and these are the males in the male/female expected(the negative exponential) if attackswere comparisonsreported below. to occur at random with respect to one another (see Three types of sampling were conductedconcur- Slater 1974). Sample size was adequate for this com- rently during each observationperiod. Throughout parisonfor only the mostfrequently attackedspecies a period ! recorded the details of all attacks by the (Purple Gallinule, Porphyrula martinica). The ob- focal pair, including time and position of attack, served and expected distributions were not signifi- identity and behavior of attacker, speciesof victim, cantly different (Kolmogorov-Smirnovtest, n = 35, and distance moved by victim during or within 5 s P > 0.20), indicating that attacksagainst this species after attack (= Aggression Sampling). Successiveat- did not tend to occur in bouts. tacks against an individual were scored as separate attacks if the attacker engaged in nonaggressivebe- havior for at least 30 s between attacks. Second, ! RESULTS recorded the behavior and position of focal pairs and their offspring every 5 min (1980) or 10 min (1981) Behaviorduring attacks.--Jacanasattack other by scan sampling (Altmann 1974) (= Behavior and birds with a combination of primarily two be- PositionSampling). Finally, ! plotted the positionsof haviors describedby Jenni and Betts (1978). I all avian intruderson the focalfemale's territory every labeled thesebehaviors "swoops" and "threats." 20 min (1980) or 30 min (1981) by scan sampling (= In the "swoop," a jacanaflies toward an intrud- Species Scan Sampling). Positions were plotted on er and strikes it with its feet, simultaneously vegetation mapsof territories. SpeciesScan Sampling uttering a sharp call. Jacanassometimes feign was momentarily interrupted to record detailsof any ongoing attacks,whereas Behavior and Position Sam- the strike by thrusting the feet toward the in- pling was completedbefore recording attack data. truder while flying past it or by veering away Observationperiods began at 0600, 0750, 0940, 1230, from the intruder just before reaching it (pers. 1350, and 1440 during 1980 and at 0600, 0900, 1200, obs.). and 1500 during 1981. Observation periods were ro- In the "threat," a iacanacrouches in front of tated randomly among male/female pairs such that an intruder and displays its carpal spurs, si- approximately equal numbers of sampleswere ac- multaneously uttering a long, shrill vocaliza- cumulatedat each time of day for each focal pair. A tion that lenni et al. (1975) labeled the "scream." total of 16 pairs was observed,but observationswere The exposed yellow spurs are highlighted concentratedon 12 pairs. Of the latter, only one in- against the maroon forewings. lacanas were dividual was observedin both yearsof the study.All but two focal animals were captured in mist-nets and never seen striking birds of other specieswith color-banded for individual recognition. their spurs.See lenni and Betts(1978) for a full Components of the above methodology, such as description of threats. observation-periodlength, varied somewhatduring On rare occasions, other attack behaviors such early 1980,but thesechanges did not biasthe results. as charges or pecks were employed. Pecks oc- To examine temporal patterns, ! divided the repro- curred as an intruder turned to flee from an ductive period of male jacanasinto four stages:Pre- attacking iacana. True fights were also rare. incubation (the period before or between nesting at- Fights,which were seenin attacksupon Purple tempts, when no eggs or offspring are present), In- Gallinules only, consistedof both participants cubation (the first day of egg-laying to the last full iumpinõ up simultaneouslyand appearing to day of incubation), Downy (day 1, the day of hatch- ing, to day 28 post-hatching,during which chicks hit each other with their feet in a brief flurry have downy plumage), and Juvenile (day 29 to dis- of activity. persal or renesting, during which offspring have ju- An attack comprised a variable number of venal plumage). The transition date between the these attack behaviors. To represent this vari- downy and juvenile stagesis somewhatarbitrary
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