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Foraging Behavior of Mangrove Swallows at Barro Colorado Island

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Foraging Behavior of Mangrove Swallows at Barro Colorado Island FORAGING BEHAVIOR OF MANGROVE SWALLOWS AT BARRO COLORADO ISLAND ROBERT E. RICKLEFS DURinG January throughJuly 1968 I had the opportunityto make incidentalstudies on a pair of MangroveSwallows (Iridoprocne albilinea) breedingin a nest box on the boathouseof the SmithsonianTropical Re- searchInstitute at Barro ColoradoIsland (BCI) in the Panama Canal Zone. Although the Mangrove Swallow is commonalong the shoresof Gatun Lake, it has receivedlittle attention. The speciesis particularly relevantto severalecological problems because it is activein openhabitats whereit is easilywatched, and becauseit has a closenorthern counterpart, the Tree Swallow(Iridoprocne bicolor). With limited time, I concentrated on nestlingdevelopment and care of the youngafter fledging(to be de- scribedelsewhere) and adult foragingbehavior during the nestlingperiod. Observationsdemonstrated complex patterns of territory utilization and diurnal activity, which are summarizedand interpretedhere. Although this report is basedon limited study of a singlepair of birds, it points up several aspectsof the use of time and spaceby birds that deservemore attention from ornithologists. GE•}m^L B•OLOG¾ The Mangrove Swallow is a rather small member (13-16 g) of the Tachycineta-Iridoprocnegroup, which includes the Tree Swallow and Violet-greenSwallow (Tachycinetathalassina) in temperateregions and severaltropical species,notably T. albiventer in northern South America and T. leucorrhoaand T. leucopygato the south (Mayr and Greenway, 1960). Three subspeciesof T. albilineaare recognized,ranging along the coastsof Central America from Sonora and Tamaulipas, Mexico, south throughPanama on both sidesof the Isthmus,and in NorthwesternPeru. Canal Zone birds belong to the nominate race. MangroveSwallows are restrictedto lowlandsincluding Gatun Lake in the Canal Zone. Like other membersof the family, the Mangrove Swallow feedsaerially, usuallyclose over lakes, larger rivers, and bays, but may forage 100 feet or more above the water surface. Occasionallyit forages over open marshesor extensivemeadows. Mangrove Swallows eat small flying insects. Henry Hespenheide analyzed the stomachcontents of two individualscollected 8 July 1968 in GuanacasteProvince, Costa Rica: one contained370 flying ants, 55 635 The Auk, 88: 635-651. July 1971 636 Ro•ar E. P,z½•c•vs [Auk, Vol. 88 July 1971] Mangrove Swallow Foraging 637 other hymenoptera,11 beetles,and 2 hemiptera; the secondcontained 13 homoptera,10 flying ants, 7 otherhymenoptera, 2 beetles,and 1 diptera. BREEDING In Panamabreeding appears to be restrictedto the dry seasonand early part of the wet season.Meager data suggestthat laying occursfrom January throughApril and that young may be fed throughJuly. Non- breedingMangrove Swallows usually forage low over the water in small groups,four or five to severaldozen, and are often seenperched on small branchesjutting from the water or on telephonewires. The onsetof the breeding seasonis marked by the breakup of flocks and presumablyco- incideswith the establishmentof nestingsites by the adults. When I left the Canal Zone for a brief trip during the third week of December1967, swallowswere still in flocks, and the rainy seasonhad not yet ended. Betweenthat time and my return to BCI 15 January, the rains had ceased, skieshad cleared,and the dry seasonwinds had pickedup. Swallowswere no longerforaging in groups,and couldbe found only in pairs around the edgeof Gatun Lake. One pair had occupieda nest box put up during the previousyear on the boathouseon BCI and was carrying nestingmaterial to it. The first eggwas laid on 19 January. The rapid changein behavior of the Mangrove Swallow at a time when changesin day length are negligible (Table 1) suggeststhat the swallowsrespond to some aspect of the abrupt wet-dry seasonchange in the Canal Zone area. Cloud cover, rainfall, relative humidity, wind velocity, foliage density in the forest canopy,and tree floweringamong other environmentalfeatures could be proximatefactors triggeringthe onsetof breedingactivity. The first flocking of swallowsafter the breedingseason in 1968 was noted on 28 May, about 3 weeksafter the young of the secondbrood at BCI fledged. Thus, the total length of the breedingseason was about 5 months. Feedingconditions evidently improvedduring the courseof the breeding season,although samples from a trap for flying insectsin a nearby clearing did not changeappreciably during this period. In Februarythe pair was able to fledgeonly oneof threeyoung in its first brood,but fledgedthree of the secondbrood, raised in April and May. Growth curves for these broods(Ricklefs, MS) showedundernourishment in the first brood but not in the second. The activitiesof the pair of MangroveSwallows at the boathouseranged over and on the peripheryof a protectedbay of Gatun Lake (Figure 1). The bay is surroundedby tall forest,except between feeding areas A and 638 Ros• E. RICKLEFS [Auk, Vol. 88 BARRO COLORADO LAKE ISLAND Figure 1. Map of the area of Barro Colorado Island and Gatun Lake showing the foraging localities of the pair of Mangrove Swallows that nested at the hoathouse. Most of the feeding occurred in areas A, B, and C. Observations were made from the edgeof the lake, about 30 m north of the nest. B, wherea clearingwith secondgrowth vegetation up to 5 m tall extends back about 100 m from the shore. The swallowsperched frequently on two buoysand a stumpin the middle of the bay, and also occasionally on the corrugatedaluminum roofs of the boathouseand a small storage shed,and on somebranches protruding from water in the area used for burning trash. The most intensive foraging areas were assignedletters and are indicatedon Figure 1 by shading,but the birds fed over the entire July 1971] Mangrove Swallow Foraging 659 Figure 2. Foraging positions of the adult Mangrove Swallows at $-10 second i.ntervals for a 12-minute period on the morning of 18 February 1969 in area A. The black circles represent feeding at heights greater than 50 feet, the lightest circles at heights lessthan 3 feet. area at times. The size of the activity area, about 500 m square,was such that the swallowscould fly acrossit anywherein 10-20 seconds. FEEDING TI-IE YOUNG Becausethe Mangrove Swallowsforaged in the open, I could record simultaneouslyboth the numberof trips to the nest and the feedingac- tivities of the adults. From one vantagepoint near the boathouseall of the swallows'feeding areas were visible except a portion of area B. I made observationsduring •6-hourperiods spaced at regularintervals when the youngof the first broodwere 13, 14, and 18 daysold, and whenthe young of the secondbrood were 4 and 13 daysold. Two youngremained in the first broodon the 13th and 14th days. The lossof weightby onenestling suggestedthat the brood was poorly nourished. On the 18th day only one young was still alive, and though it was losing weight at this time, re- cessionof weight is normal for older nestlingswallows (Ricklefs, 1968) and its weight was greater than that of adults. The secondbrood con- tained three well-nourishedyoung during the entire nest period. On the 4th day energy requirementsof the young would have been low owing to their small size. I also watched the swallowsduring the egg period of the second brood. 640 ROBERTE. RICKLEFS [Auk, Vol. 88 TABLE 2 FORAGING BEI-IAVIOR OF ADIJLT MANGROVE SWALLOWS ON BARROW COLORADO ISLAND FEBRUARY 1968 Time spent Time spent out Time of Number of flying of nest area observation feedings (per cent) (per cent) 18 February• 06:30-07:00 28 94.8 5.0 08:30-09:00 7 53.9 11.8 10:30-11:00 8 29.4 46.7 12:30-13:00 17 42.4 12.3 14:30-15:00 6 34.1 75.6 16:30-17:00 1 86.7 28.1 19 February• 06: 40-07: 00 11 93.8 19.0 07:30-08:00 12 46.2 17.4 09:30-10:00 10 47.3 41.5 11: 30-12:00 15 53.1 14.1 13: 30-14: 00 8 56.6 20.0 15:30-16:00 3 80.9 51.2 17:30-18:00 10 79.7 5.0 23 February• 06:30-07:00 7 91.8 24.1 07:30-08:00 4 33.7 16.7 08:30-09:00 12 37.5 18.0 09:30-10:00 10 65.9 49.4 10:30-1i:00 1 15.0 13.3 11:30-12:00 7 33.6 1.3 12:30-13:00 9 52.1 9.0 13:30-14:00 8 38.5 8.7 14:30-15:00 7 45.5 4.1 15:30-16:00 5 47.9 0.8 16:30-17:00 7 35.3 4.7 17:30-18:00 4 67.8 13.7 Two nestlings 13 and 14 days old. One nestling 18 days old. Adult swallowsused two distinct foragingpatterns while feedingyoung in the nest box. The first consistedof foraging within the immediate vicinity of the nest, usually within 100 m and rarely beyond 150 m, roughlyarea A on Figure 1. The young were fed frequentlyduring these periodsof activity. During one period the positionsof the adults were recorded every 5 to 10 secondsfor 12 minutes (07:10-07:22) on the morningof 18 February, the 13th day of the nestlingperiod (Figure 2). At this time, most of the foragingoccurred 5-15 m abovethe water, and a large portion of the activity centeredabout the small inlet around the dock and boathouse. At other times the feeding pattern extended farther out towardsthe bay in front of the boathouse,closer to the water surface, or perhapsover the trees at the edgeof the water. The adult swallowsspent periodsvarying from as few as 1-2 minutes July 1971] Mangrove Swallow Foraging 641 6 8 I0 NOON 2 4 6 • I00- • 6 8 I0 NOON 2 4 6 50 o • o [-1 I-] m • 6 8 I0 NOON 2 4 6 TIME OF DAY Figure 3. Foragingcharacteristics of the pair of MangroveSwallows on 18 and 19 February when two well-grown nestlingswere present.
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