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Polygyny in the Asian Openbill (Anastomus Oscitans)

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Polygyny in the Asian Openbill (Anastomus Oscitans) January1995] ShortCommunications and Commentaries 257 steroid measurementsto the assessmentof gonadal faecal steroid analysis.J. Zool., Lond. 224:649- activity in free-living Kakapo. The survival of this 668. endangered speciesdepends on egg laying by female COCKP,EM, J. F. 1989. Reproductivephysiology and Kakapo. Thus, conservation efforts are focussedon the managementof the Kakapo.Ecology Division the females. The shortageof female samplesin our Report No. 18. Lower Hutt, Ecology Division, study meansthat further samplecollection programs Department of Scientificand Industrial Research, are neededto realize the potential of this new method Lower Hutt, New Zealand. to provide information on the reproductivecondition COCKREM,J. F., AND J. R. ROUNCE. 1994. Faecal mea- of female Kakapo. In a wider context, fecal steroid surements of estradiol and testosterone allow the measurementscan now be appliedto otherfree-living non-invasive estimation of plasma steroid levels birds for which handling and blood sampling is dif- in the chicken. Br. Poult. Sci. 35:433-443. ficult or inappropriate. FOLLETT,B. K. 1984. Birds. Pages283-350 in Mar- Acknowledgments.--Wethank T. Greene, B. D. Lloyd, shall's physiology of reproduction,4th ed. Vol. R. G. Powlesland,C. Smuts-Kennedy,and other work- 1, Reproductivecycles of vertebrates(G. E. Lam- ers on Little Barrier Island for the collectionof drop- ming, Ed.). Churchill Livingstone,Edinburgh. pings.B. D. Lloyd, D. V. Merton, and R. G. Powlesland KOFUJI,H., M. KANDA,AND T. OISHI. 1993. Breeding generously provided information about Kakapo. S. cyclesand fecal gonadal steroidsin the Brown Ishii and C. M. Bishopkindly made available details Dipper Cincluspallasii. Gen. Comp. Endocrinol. of their methodsfor fecal-samplepreparation. Tl•e 91:216-223. work was funded by the Threatened SpeciesTrust POWLESLAND,R. G., B. D. LLOYD, H. A. BEST,AND D. through the Department of Conservationas part of V. MERTON. 1992. Breeding biology of the Ka- the Kakapo RecoveryProgramme, and D. J. Melior kapo Strigopshabroptilus on Stewart Island, New provided accessto laboratoryfacilities. Zealand. Ibis 134:361-373. WILKINSON,L. 1988a. SYGRAPH. Systat,Evanston, LITERATURE CITED Illinois. WILKINSON,L. 1988b. SYSTAT. Systat,Evanston, Il- BERCOVITZ,A. B., J. COLLINS,P. PRICE,AND D. TUTTLE. linois. 1982. Noninvasive assessment of seasonal hor- WINGFIELD,J. C., G. F. BALL,A.M. DUFTY,JR., R. E. mone profiles in captive Bald Eagles(Haliaeetus HEGNER, AND g. RAMENOFSKY. 1987. Testoster- leucocephalus).Zoo Biol. 1:111-117. one and aggressionin birds. Am. Sci. 75:602-608. BISHOP,C. M., AND M. R. HALL. 1991. Non-invasive monitoring of avian reproduction by simplified Received15 September1993, accepted 12 June1994. TheAuk 112(1):257-260, 1995 Polygyny in the Asian Openbill (Anastomusoscitans) TANMAY DATTA AND B.C. PAL Departmentof Zoology,North BengalUniversity 734430, Dist. Darjeeling, West Bengal, India Although monogamy is the predominant mating We describeseveral cases of polygyny in a breeding systemin birds, there are few strictly monogamous population of a stork, the Asian Openbill (Anastomus bird species(see review by Ford 1983). Other than oscitans).We also report information concerningthe the widespread occurrenceof polygyny in various formation of suchmating groupsand the successex- species,occasional polygyny also has been reported perienced by them in comparison to monogamous in normally monogamousspecies (Armstrong 1955, pairs. Verner and Willson1969, Logan and Rulli 1981,Marks Methods.--Our study was conductedat the Raiganj et al. 1989). Although Brown (1987) did not include Wildlife Sanctuary(25ø36'N, 38ø10'E),West Bengal, any membersof the Ciconiiformesin his list of 222 India, where AsianOpenbills are found breedingfrom speciesof communallybreeding birds, occasionalpo- July through December,as are five other waterbird lygyny also has been found in this group (Lancaster species. 1970, Cramp 1977, Fujioka 1986, McKilligan and We studied this population of openbills from 1987 McConnell 1989).However, storksalways have been to 1991. Nests with more than two adult birds were found to breed monogamously(Ali and Ripley 1968, consideredas a possible caseof polygyny, and we Cramp 1977, Coulter et al. 1989). kept closewatch on thosenests. Data from thosenests 258 Short Communicationsand Commentaries [Auk, Vol. 112 were taken into accountonly when the presenceof rushed to that spot and got involved in the distur- more than two adult birds was confirmed by direct bance.In these cases,fighting continued for 5 to 18 observation for at least three consecutivedays. min (2 = 588 + SD of 296.74 s), involving 6 to 13 Membersof 37 suchgroups were colormarked with birds (2 = 9.29 _+2.29). The duration of fighting was nontoxic textile dyes using a sprayer from a high directly proportional (r = 0.946) to number of birds vantage position above the nest-rim level. For each involved. At the end of fighting, three or four birds nest we maintainedan identity card with the records were found to rest peacefullyin that nesting spot. of the position of the nest in the sanctuaryand the The new group sometimesincluded one or both of marking patternsor identifiable physicaltraits of the the original pair, but this was not always the case. members. Sexes were ascertained from behavior. Po- Within a very shortperiod, the birdsat the nestmade sition during copulationwas treated as a good indi- joint up-down displays(Kahl 1972) in very closesuc- cator of sex, as "reverse mounting" is very rare in cession.Thereafter, they behaved as a unit in all re- this species.Above all, we think our continuousand spects,exhibiting pair-forming displays,as done by careful observationswere effective in making certain monogamouspairs. Such intraspecificattacks were that we did not introduce the confounding effect of very commonin this colony and, in most cases,they reverse mounting. ended with only two birds in the nest. Nestswere inspectedeach day beforehatching and We were able to surveythe developmentand suc- thereafter on alternate days. Observationwas made cessof clutchesat 11 polygamousnests. Polygamous from a high vantageposition or from the groundwith clutchesshowed greater variation in layingorder than minimum interference to birds in the area. To com- those in monogamousnests, where eggs were con- pare successrates, four monogamouspairs were se- sistentlylaid on alternatedays. In polygamousnests, lectedas controlsfor eachpolygamous nest. Control mostof the eggswere laid on alternatedays; however, nestswere selectedfrom the closevicinity of the po- in six instanceseggs were added to the clutch on the lygamousnest; the physicaland breedingstatuses of sameday. In another nest, there was a three-day in- all control birds were similar. Observations were made terval between eggs.Clutch size of polygamousnests with the naked eye or with 20 x binoculars. Mea- (œ= 4.45 _+ 1.44, n = 11) was significantly higher (t surementsof eggswere taken with a vernier caliper, = 3.76, df = 53, P < 0.001) than that for the control and massesof young were taken with a Pesolaspring monogamousnests (2 = 3.41 _+0.54, n = 44). balance. Intraclutch variation of egg dimensiondid not fol- Results.--Duringthis five-year study, we found 51 low definite order in polygamousclutches, in contrast monogamousgroups of 2,115 active nests.In 2 cases, to the situation in monogamousclutches. In monog- four birds used a single nest, and the other 49 nests amousclutches, larger eggsalways were laid in the were occupiedby trios. middle of the clutch.In three- or four-egg clutches, In this colony, openbills usually started nesting the secondegg was largest,while in five-eggclutches from the first week of July. The polygamousnesting the third was largest.In polygamousclutches, no such attemptswere not seenuntil August,when almostall order was maintained and, in some cases, the first or of the nesting trees were invaded by monogamous last egg of the clutch was the largest.Intraclutch dif- pairs. Nesting by monogamouspairs also continued ferencesbetween the largest and smallestegg were until the lastweek of September.After the third week significantlygreater in polygamousnests (t = 3.76, df of July, most of the new settlershad no option other = 53, P < 0.001). In 44 control monogamousclutches, than to choosea preoccupiednesting tree for nesting, the range of egg volumes was never more than 30% which often resulted in intraspecificfighting. Natu- (œ= 14.8%)of the smallestegg volume. However, in rally polygamousgroups always nested in a preoc- 8 of 11 polygamousclutches, this differencewas more cupied nesting tree with many monogamousnests. than 35% 07 = 30.6%). Polygamousnests were not confinedto any particular Hatching successand fledging successwere con- zone of the colony, but were scatteredthroughout. siderablyhigher in polygamousnests, while lossdue Copulatory behavior confirmed the sexesof the to intraspecificattack was higher in monogamousnests members of 21 trios in which two females mated with (Table 1). The massof young birds at fledging was a common male. In 15 other cases,behavioral (other significantly higher (t = 3.2, df = 49, P < 0.005) in than copulation)and physicalcharacters of the mem- polygamousnests. bersalso suggested that only one male wasinvolved. All members of the polygamous nests shared in However, for the remaining 15 cases,sexes of the mostof the parentalactivities, from nest building to membersof thosemating groupswere not identified. food provisioningof young. We sawno obvioussigns
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