Nest Attendance of Parent Birds in the Painted Snipe (Rostratula Benghalensis)

NEST ATTENDANCE OF PARENT BIRDS IN THE PAINTED SNIPE (ROSTRATULA BENGHALENSIS)

SHIGEMOTO KOMEDA Laboratoryof AnimalSociology, Faculty of Science,Osaka City University, Sugimoto,Sumiyoshi-ku, Osaka 558, Japan

ABSTRACT.--Iobserved 15 pairs of Painted Snipes (Rostratulabenghalensis) in Japanfor 24-h periods in order to compare the nest attendance behavior of males and females and to documentchanges in their behavior during the laying period. From a few days before the laying of the first egg until the day the secondegg was laid, the male and female consorted with eachother (stayedwithin 5 m of each other) more than 90% of the observationtime. They visited the nest repeatedlyon the first and seconddays of laying. On the third day, the consortingratio decreasedto 58.5%, and the male typically started to incubate. He continuedto incubate that night. After laying the third egg, the female usually did not visit the nest exceptto lay the fourth and last egg. The male continuedincubation, and the female met her mate only when she laid the last egg. The absenceof female incubationand the early termination of consortingbehavior are consistentwith the possibility that Painted Snipe femalesare polyandrous.Received 12 January1982, accepted4 October1982.

AMONG the polyandrous charadriiform qualis), and cudweed (Gnaphaliumaffine), grew species, comprehensive studies of pair-bond sparselyin theseabandoned fields. The water depth maintenancehave been conductedon only four in this area varied from 0 to 15 cm. species, the Northern Jacana(Jacana spinosa; The main study was conducted in Toyo City, northwestern Shikoku Island, Japan (133ø03'E, Jenni and Collier 1972, Jenni and Betts 1978), 33ø57'N),from March to August1979 (excluding May) the SpottedSandpiper (Actitisrnacularia; Hays and from June to July 1980 (Toyo area). This 40-ha 1972, Oring and Knudson 1972, Maxson and study area of reclaimedland facesthe JapaneseIn- Oring 1980),the Red-neckedPhalarope (Phal- land Sea. A quarter of the area was used as paddy aropuslobatus; Hild6n and Vuolanto 1972),and fields, and the rest was marshy grasslandthat was the Red Phalarope(Phalaropus fulicaria; Scha- plowed onceor twice a year, usuallyin Juneand July. mel and Tracy 1977).Although a detailedstudy The study area was traversedby six parallel roads of the Painted Snipe (Rostratulabenghalensis) 200m apartand by anotherroad that intersected them has not been conducted, sex role reversal and at right angles.This poorly drained areawas always an apparentlyunbalanced sex ratio suggestthe covered with water. Until it was plowed, dense reed (Phragmitescommunis) or bulrush (Scirpusplanicul- possibility of polyandry in this species(Beven mis) grew there every year. After it was plowed, the 1913, Baker 1935, Lack 1968, Hays 1972, Oring grassesgrew only patchilyor sparselybefore Sep- and Knudson 1972,Jenni 1974).The objectives tember. The Toyo area is about 300 km west of the of this study were (1) to compare the nest at- Biwa area. tendance of male and female Painted Snipes, Most data for the field study were obtained by con- (2) to measurethe amount of time the members tinuous observations. Observations that started after of the pair spend together (consortingratio) 1800 are identified as "nocturnal observations" and during laying and incubationperiods, and (3) were made with the aid of a flashlight.Behavior was to considerthe evidence for polyandry. observedwith binoculars(9x) and a spotting scope (15x). Behavioral actswere recorded in seconds,and the data were later converted to minutes for calcu- STUDY AREAS AND METHODS lations. Of 27 birds observed, 4 males and 2 females Field observations were made in two areas. The were marked with neck bands. Because no other birds preliminary study was done in Chuzu Town, south- of the same sexes as the banded individuals were east of Lake Biwa, central Japan(136ø02'E, 35ø08'N), observedentering their nests, any male or female from May to June1978 (Biwa area). Nest observations who entered a nest was consideredthe progenitor of were conductedin 20-ha abandonedfields among the eggsin the nest. paddy fields at an elevation of about 85 m. Marshy All observationsreported here were made at nests plants, mainly buttercup(Ranunculus quelpaertensis), with four eggs.The four eggswere laid on consec- bitter cress(Cardamine lyrata), foxtail (Alopecurusae- utive days,and the daysof layingwere identifiedas 48 The Auk 100: 48-55. January1983 January1983] NestAttendance in Painted Snipes 49

T^BLE1. The lengthof visitsto the nestduring the layingperiod (minutes + SD).Numbers in parentheses indicate nocturnal visits.

Day I Day 2 Day 3 Day 4

Female Lengthof egg-layingvisits 35.5 + 17.5 62.3 + 33.4 32.8 + 7.4 15.9 + 1.9 Number of egg-layingvisits 2 4 5 6 Length of other visits 3.6 + 3.7 6.1 + 5.2 5.7 + 5.3 I -- Number of other visits 43 50 36 I

Male Length of visits 3.6 + 8.0 8.0 + 12.8 22.6 + 31.7 43.4 + 52.8 (21.6 + 15.1) (36.1 + 10.2) Percentageof stayslonger than 2.0 6.8 26.2 46.8 30 min (35.3) (76.9) Number of visits 99 85 92 47 (17) (13) Number of nests 4 7 9 8 (1) (1)

day 1, day 2, and so on. Thesefour dayscomprise During the laying period, a female laid an egg the layingperiod. A spanof a few daysbefore day every morning between 0700 and 1100(Fig. 1). I is referred to as the prelaying period. The incu- She stayedon the nestfar longerwhen coming bationperiod extends from the day afterday 4 until to lay an eggthan at other times(Table 1). The hatching. female was on the nest for a much shorter du- The percentageof time during which the two ration when she laid the fourth egg (2 = 15.7 membersof a pair stayedwithin 5 m of eachother was termedthe consortingratio. Consortingbehav- min) than when she laid the earlier eggs (2 = ior was studiedat the Toyo areafor a total of 17 days 44.0 min). All other visits to the nest by the (162.0h). Other observationswere madeat both the female during the laying period were much Toyo and Biwa areas. shorter(2 = 5.1 min); 88.5% of thesevisits were Nest-related behavior in the Biwa area was ob- less than 10 min. served from blinds about 10 m from the nests. Four On four additional days when the 1st, 2nd, nestswere observedfor a total of 17 days (190.8 h). or 3rd eggs were laid but egg laying was not Observationswere made at the Biwa area during two observed,females stayed on the nest for 39, 42, nights(22.6 h) of the incubationperiod. All diurnal 46, and 47 min between 0800 and 1210 (Fig. 1; observationsat the Toyoarea were made from a blind 22 June 1979, 21 June 1978, 3 June 1978, and 12 on a carroof 1.5 m abovethe ground.By driving the car alongthe roads,I was able to follow the birds June 1979, respectively).These four stayswere whenthey left their nests.Nine pairswere observed the only long stay on thosedays. It is assumed at the Toyoarea for a total of 36 days(368.7 h) Two that egg laying occurredduring theselong vis- of thesenine pairsabandoned their nestsduring the its. Egg laying was not seen at two nests on layingperiod, so the datafrom thesenests were ex- day 4, but the females entered the nests only cluded from the results(total 7 days).Two nestswere once, and it is assumed that they laid during observeda total of four nights (47.0 h) during the thesevisits (Fig. 1; 30 March 1979 and 21 July layingperiod. Activity at night at the Toyoarea was 1979). observed at intervals of 1 min from about 20 m when Nestvisitation on eachlaying day.•On day 1, a flashlightwas switched on for about5 s and off for about 55 s. both sexesentered the nest frequently during the morning (Fig. 1), but they often left it for

RESULTS a long time during the afternoon. They re- turned to the nest in the evening. The male Clutch size and time of laying.•Of 33 nests stayedabout the samelength of time (Table 1) found, 2 had 3-egg clutches,30 held 4 eggs, and enteredthe nest about as frequentlyas the and only 1 held 5 eggs.Accordingly, the nor- female did. No observations were made on the mal clutch size in this speciesis consideredto night of day 1. be four. On day 2, the duration of diurnal visits in- The time of laying was recorded 17 times. creased but the number of visits decreased 50 $HIGEMOTOKOMEDA [Auk,Vol. 100

TIME OF DAY DAYTIME ACTIVITY 6 9 12 15 18 DAY DATE NEST ,

I 12• JUN '78'79 62 2 AUG ' 79 9 9 .N.JG '79 I0 4JUN '78 2 15 JUN '78 :• 20 JUN '78 4 28 MAR ' 79 5 13 JUN ' 79 6 22 JUN '79 7 3' AUG ' 79 9 I5 MAY '78 I 5 JUN '78 2 16 JUN '78 3 21 JUN '78 4 29 MAR '79 5 I 4 JUN ' 79 6

23, JUN '79 7

4 AUG '79 9 I I AUG '79 I I 16MAY '78 I 6 JUN '78 :• 17 JUN '78 '• 22 JUN '78 4 :30 MAR '79 5 2[ JUL '79 8 5 AUG '79 9 12 AUG '79 I I

NIGHTTIME ACTIVITY 18 21 0 3 6 DAY DATE NEST 2 i8-19 JUN'80 14 2 24-25JUL '80 15 • 25-26JUL '80 15 4 26-27JUL '80 15

Fig. 1. Activity during laying period. Male is shown in upper and female in lower belt of each pair. Blackband shows time spenton nestand shadedband out of nest.Small dot shows the time of egglaying. Day 1 is the first day of egglaying and day 4 the last day. slightly in both sexes.In two nestswatched at returned at 0443. At Nest 14 the male and fe- night, both sexes left the nest site just after male were gone when observationsstarted at dark and returnedjust beforelight. At Nest 15 2100 and returned to the vicinity (within 5 m) the male and female left the nest at 1907 and of the nest at 0426. January1983] NestAttendance in PaintedSnipes 51

TABLE2. The percentageof daylight hours during N,,4 which the male and female of a pair stayedwithin 5 m of each other. t N,,7 Number of Num- hours ber I N-8 ob- of Average (range) served pairs a'o 4'0 do 8b o'o Prelaying % OF TIME period 99.6 (98.5-100.0) 23.6 4 Fig. 2. Average percentageof daylight time on Laying period nest. Upper belt shows male stays and lower belt Day 1 99.0 (97.5-100.0) 34.0 3 female stays. Blackband shows percentageof time Day 2 92.8 (89.3-95.9) 42.6 4 spent on nest and white band out of nest. Number Day 3 58.8 (38.4-81.0) 29.7 3 of nests observed is shown. Day 4 7.9 (3.3-16.7) 32.1 3

On day 3, both sexeschanged their patterns day 4 she stayedon the nest much longer than of nest visitations, but in different ways. Fe- the other females. On her first visit she stayed malesentered the nestrepeatedly before laying for 27 min and laid the fourth egg. She then the third egg but after that did not return to flew off and returned 80 min later and stayed the nest until 1800 in all but two nests. Females on the nest for 5.5 h. Neither the male nor the entered three of four nests between 1800 and female returned after day 4. 1930. At Nest 15 the female entered the nest at Total time spent on nest in laying period.- 1851,left at 1903,and did not return that night, During the whole laying period of daylight but her mate spentthe night on the nest. Dur- hours, males stayed on the nest four times as ing the day malesvisited the nestabout as often much as females(39.8% vs. 9.9%). As egg lay- as the previousday, but the averagevisit was ing advanced,males increasedand femalesde- much longer (Table 1). creasedthe amount of time they spent on the On day 4, all but one of the females entered nest (Fig. 2). On days 1 and 2, females spent the nest only onceand laid the last egg at that slightly more time on the nest than males did time. After leaving the nest, they were never (16.4% vs. 13.2% and 16.7% vs. 16.2%, re- seen near it again. At Nest 9 the female left the spectively).On day 3 somemales spent a long- nest i min after entering it, but she returned er time on the nest than previously, and it was 30 s later and laid the fourth egg.Males entered on this day that the greatestrange was seen in the nest less often than previously but stayed the percentagesof time spent on the nest for muchlonger during each visit. The malesstayed different males. The males on five of the nine on the nestat night, but femalesdid not return nests spent more than 50% of the day on the to the nest. nest (46.2% on average,range 16.3%-71.3%). Two males in the Toyo area abandonedtheir The femaleson day 3 reducedtheir time on the nests after their femaleslaid the third egg, but nest to only 7.3%. On day 4 the males spent the femalesstill laid the fourth eggin the nests. 78.7% of the day on the nest, and femalesspent Pair 12 visited their nest in typical fashion only 3.0%. through day 3. On day 4, the female exhibited Femalesdid not visit their nestsat all at night typical behavior. She enteredthe nest, laid the during the laying period, and males did not fourth egg, left the nest, and never returned. visit at night until after the third egg was laid. The male on day 4, however, never visited the A male spent68.6% of the night followingthe nest, nor was he seenaround the nest. I began laying of the third egg and 86.7% of the night observingNest 13 on day 2. The male did not following the laying of the fourth egg on the enter the nest between 0744 and 1157 on day nest. 2, nor at any time on day 3, and he did not Male nest attendance in the Painted Snipe even visit the vicinity of the nest on day 4. His reached mid-incubation levels on day 3, and female stayed on the nest more than 40% of the incubation started on day 4. In the Red- the total observation time on both days. On necked Phalarope (Hild6n and Vuolanto 1972), 52 SHIGEMOTOKOMEDA [Auk, Vol. 100

DAY I DAY 2

40-

II I- • 60-

DAY 4 40-

20-

o 0-5I 5-10 DAY10-1515-20 20< 0-5 5-10 10-15 151-20 DISTANCE FROM NEST (m)

Fig. 3. Average distanceof female from nest. Shaded bar shows time female spent with male and white bar shows time female spent alone. male visits to the nest become gradually more Because the male and female arrived at the frequent and longer as the laying period pro- nest togetherearly in the morningbefore the gresses.The SpottedSandpiper male incubates laying of the third egg, it is possiblethat they sporadicallybefore day 3 (Maxsonand Oring had consortedduring the precedingnight. The 1980).In the Red Phalarope(Schamel and Tracy female probably left her mate on the nest at 1977, Mayfield 1979), the Wilson'sPhalarope night after the third egg was laid, however, (Phalaropustricolor; Howe 1975), and the Wat- becausethe male began spending considerable tled Jacana(Jacana jacana; Osbone and Bourne time on the nest and the femaledid not stay in 1977), males appear to begin incubating on the vicinity of the nest. day 3. As egg laying advanced,the amount of day- Consortingratio.--A male usuallyfollowed a light time femalesstayed within 5 m of the nest female for three or four days during the pre- decreasedin the Toyo area (Fig. 3). On days 1 layingperiod. The consorting ratio was 99% in and 2, femalesstayed within 5 m of the nest the Toyo areaduring the prelayingperiod. Al- most of the time and consorted with the males though the consortingratio for the whole lay- almostall day. Within 10 m of the nest, females ing periodwas 68%, it decreasedfrom 99% on were occasionallyseparated from the males, day 1 and 93% on day 2 to only8% on the last who stayedon the nest, but when femaleswent day (Table 2). The consortingratios varied more than 15 m from the nest, the malesalways widely among individual pairs on day 3. In followed. On day 3 females spent less time two of the threepairs the femaleconsorted with within 5 m of the nest and often went away the male on day 4 only when she came to lay. from the nest alone. After laying the fourth egg, One female (Nest 9), however, returned to the femalesleft the nest and movedsolitarily, while vicinity of the nest after laying the fourth egg the males stayedon the nest. and stayedwithin 5 m of the incubatingmale. Behaviorduring the incubationperiod.--Dur- January1983] NestAttendance in Painted Snipes 53

TABLE3. Nestattendance of maleduring the incubationperiod. (Females did notvisit the nestduring the incubation period.)

Daytime Nighttime Averagepercentage time on nest(range) 80.4(64.9-94.7) 89.6(88.2-91.1) Averageduration of nestvisits in minutes(range) 39.8(23.9-110.4) 92.9(77.7-111.2) Percentageof visitsexceeding 30 min 53.4 100.0 Numberof days 14 2 Number of hours observed 156.0 22.6

ing the incubationperiod males typically left other hand, only the males incubate in the the nest 1.3 times per hour for about 10 min. Northern Jacana(Jenni and Betts 1978), the Red- Males sometimes remained on the nest for sev- neckedPhalarope (Hild6n and Vuolanto 1972), eral hours, especiallyduring the morning. At and the Red Phalarope(Mayfield 1979), all of othertimes, they stayed away froin the nestfor which have proved to be polyandrous.In the aslong as 40 min. Maleswere on thenest 80% Wilson's Phalarope (Howe 1975), which is of the time during the day and 89% during the probably polyandrous,only males incubate. night (Table3). At night the malesleft the nest The Painted Snipe female decreasedthe time lessfrequently and spentlonger periods on the she spenton the nest as egg laying advanced, nest. and almost all her nest visits occurred before When malesleft the nest,they usuallywalked the third egg was laid and were less than 10 or flew away immediately.In 61 of 85 obser- min long, exceptfor the four egg-layingvisits. vations made during the incubation period, After a male started to incubate, the female did males left the vicinity of the nest within 1 min not visit the nest againexcept to lay the fourth of leaving the nest. This behavior differed egg, nor was she ever seenin the vicinity of markedly froin behavior during the laying pe- the nest. Therefore, incubation is performed riod, becausethe male spent much time in the only by a male.A femaleneither helps her mate vicinity of the nest during the laying period. nor incubatesalone. This suggeststhat the de- A male typicallyflew back to a placeabout 20 gree of sex-rolereversal in incubationis more in froin the nest and then walked to the nest, complete in the Painted Snipe than in the where he preened for a time before entering Mountain Plover, the Sanderling, or the Spot- the nest. ted Sandpiper and is similar to that in the The incubation period averaged 16.8 days Northern Jacanaand the three speciesof phal- (range 16-18 days, n = 6). During the incu- aropes. One Painted Snipe female began in- bationperiod, the femalewas neverseen at the cubatingwhen her mate abandonedtheir nest nest and apparentlynever cameanywhere near but ceasedafter about a half-day.This fact may the nest (confirmedby banded femalesfor 7 suggestthat ancestralPainted Snipe females days). When the male interruptedincubation incubated. to forage,he did not consortwith any females In the speciesin which femalesdo not in- (confirmedfor 6 days).The maleand his chicks cubate,females may deserttheir mates during left the nest within a half-day after all the eggs incubation, as in the Red Phalarope (Schamel hatched,and they never returned to their nest and Tracy 1977, Ridley 1980); they may help (n = 4). their mates, as in the Northern Jacana, the fe- males of which visit the nest, shade the eggs DISCUSSION (Jenniand Betts 1978), and help defend the ter- In speciessuch as the MountainPlover (Cha- ritory againstother birds, even during the in- radrius montanus;Graul 1973) and the Sander- cubationand rearingperiods (Jenni and Collier ling (Calidrisalba; Panneleeand Payne 1973), 1972,Jenni 1974); or they may deserttheir mates which have double clutches, females incubate as soon as or before the clutch is completed, as the last clutch with or without males. In the in the Red-necked Phalarope (Hild6n and Vuo- polyandrousSpotted Sandpiper, females help lanto 1972) and the Wilson's Phalarope (Howe with incubation unlessthey obtain a new mate 1975). In the Painted Snipe, males did not fol- (Hays 1972,Maxson and Oring 1980).On the low their mates on or after day 4, and females 54 SHIGEMOTOKOMEDA [Auk, Vol. 100

did not visit their nestsafter the fourth eggwas She copulatedonly with the later mate during laid. The pair bond of the Painted Snipe ap- the layingperiod of the earliermate (Oring and peared to break down after the laying of the Maxon 1978). In these cases, the earlier male last egg in the same way as the pair bonds of does not always rear his own offspring, al- Red-necked and Wilson's phalaropesdo. It is though Northern Jacana females spend a advantageousto a sequentiallypolyandrous fe- greateramount of time in the earlier male's ter- male that she desert the first mate as soon as ritory just beforeand during the laying period possibleand gather energy reservessufficient and simultaneousmating is exceptionalin the to obtain the next mate and to lay his clutch. Spotted Sandpiper. Other mechanismsmay In the simultaneouslypolyandrous American explain fertilization in specieswith a simulta- Jacana,a female stays with her mate and pre- neously polyandrousmating system.Physio- pares to lay the replacement clutch for him logical and genetic researchon these species (Jenni and Collier 1972). will be necessaryto identify these mecha- Painted Snipe females are unlikely to mate nisms. with anothermale during the prelayingperiod or on the first and seconddays of laying, be- ACKNOWLEDGMENTS causethey consortwith their currentmates most I thank Prof. Donald A. Jenni of the University of of the time on thesedays. Femalesalso consort Montana and Ms. Helen Hays of the American Mu- with their matesduring the night after the sec- seumof Natural History for their criticalreading and ond egg has been laid. During day 3, females revision of the manuscript.I wish to thank the mem- begin spending less time with their mates, but bersof the Laboratoryof Animal Sociology,Faculty by this time it is likely that their fourth egghas of Science,Osaka City University,especially Dr. Sa- alreadybeen fertilized. It is extremelyunlikely toshi Yamagishi, for continuing guidance and en- that even the last egg could be fertilized by couragement.I alsowish to thank Dr. NagahisaKu- roda of YamashinaInstitute for Ornithologyfor his some other male. After laying the last egg and instructive comments. abandoningtheir mates, the femalescould be- gin to consortwith other males. The abandon- LITERATURE CITED ment of the mate and emancipation from in- B,•I(ER, E. C.S. 1935. Nidification of birds of the cubation make it possible for females to mate Indian Empire. London, Taylor and Francis. with other males, thus enabling this speciesto BEVEN, J. O. 1913. Notes and observations on the have a sequentiallypolyandrous mating sys- PaintedSnipe (Rostratulacapensis) in Ceylon.Ibis tem. 54: 527-534. A simultaneously polyandrous female cop- GR•,UL,W.D. 1973. Adaptive aspectsof the Moun- ulates with several males in a short time. In tain Ploversocial system. Living Bird 12: 69-94. contrast, a sequentially polyandrous female Hays, H. 1972. Polyandryin the SpottedSandpip- mates with only one male during a set period. er. Living Bird 11: 43-57. The probability of a male rearing his own off- HILD•N, O., & S. VUOLANTO. 1972. Breeding bi- spring is different in these two polyandrous ology of the Red-neckedPhalarope Phalaropus lobatus in Finland. Ornis Fennica 49: 57-85. sytems. In the sequentially polyandrous HOWE,M. A. 1975. Behavioralaspects of the pair species,a male has only to keep his mate for a bond in Wilson'sPhalarope. Wilson Bull. 87: 248- few days before and during the laying period 270. to ensure that he will rear his own offspring JENm,D.A. 1974. Evolutionof polyandryin birds. (Schamel and Tracy 1977). A simultaneously Amer. Zool. 14: 129-144. polyandrousfemale lays the first set of eggson --, & B. J. BETTS. 1978. Sex differences in nest one nest for the first mate and the second set construction, incubation, and parental behav- of eggs on another nest for the second mate, iour in the polyandrousAmerican Jacana (lacana successively(Jenni and Collier 1972, Maxson spinosa).Anim. Behav. 26: 207-218. and Oring 1980),but she copulateswith several ß & G. COLLIER. 1972. Polyandry in the American Jacana(lacana spinosa).Auk 89: 743- males during that period. Northern Jacanafe- 765. males copulatedwith three or four different L,•ci•, D. 1968. Ecologicaladaptations for breeding males in a single afternoon (Jenni and Collier in birds. London, Methuen & Co. 1972). One Spotted Sandpiper female copulat- MAXSON,S. J., & L. W. ORING. 1980. Breedingsea- ed with two different males for two days, and sontime and energybudgets of the polyandrous she laid for the earlier mate on the next day. Spotted Sandpiper. Behaviour 74: 200-263. January1983] NestAttendance in PaintedSnipes 55

MAYFIELD,H. F, 1979. Red Phalaropesbreeding PARMELEE, D. F., & R. B. PAYNE. 1973. On mul- on BathurstIsland. Living Bird 17: 7-39. tiple broodsand the breedingstrategy of arctic ORING,L. W., & M. L. KNUDSON.1972. Monogamy Sanderlings.Ibis 115: 218-226. and polyandryin the SpottedSandpiper. Living RIDLEY,M.W. 1980. The breedingbehaviour and Bird 11: 59-73. feeding ecologyof Grey PhalaropesPhalaropus --, & S. J. MAXSON. 1978. Instances of simul- fulicariusin Svalbard.Ibis 122:210-226. taneouspolyandry by a SpottedSandpiper Ac- SCHAMEL,D., & D. TRACY. 1977. Polyandry, re- tiris rnacularia. Ibis 120: 349-353. placementclutches, and site tenacity in the Red OSBORNE,D. R., & G. R. BOURNE. 1977. Breeding Phalarope (Phalaropusfulicarius) at Barrow, behavior and food habits of the Wattled Jacana. Alaska. Bird-Banding 48: 314-324. Condor 79: 98-105.

IMPORTANT NOTICE TO AUTHORS

Effectivewith this issue (January1983), The Auk will adhere to use of the commonand scientificnames of North American birds as listed in the Thirty-fourth'Supplement to the American Ornithologists' Union Check-List of North American Birds (1982 Auk 99(3): 1CC-16CC). Please use this nomenclaturewhen pre- paring manuscriptsfor submissionto TheAuk. Exceptionsshould be justified in writing to the Editor.