Colony and Nest Site Selection in White-Faced and Glossy Ibises
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COLONY AND NEST SITE SELECTION IN WHITE-FACED AND GLOSSY IBISES JOANNA BURGERAND LYNNE M. MILLER ABSTRACT.--The factors determining colony and nest site selectionin the White-faced and GlossyIbises (Plegadis) were examinedfrom 1972through 1975. White-faced Ibises were studied in Texas and Argentina, GlossyIbises in New Jerseyand New York. White-faced Ibises generally nest in tule marshes,infrequently on dry land. In selectinga dry land site, White-faced Ibises preferredto nest on the ground among low shrubsand in mixed forbs rather than in habitats containinggrass and cactus. GlossyIbises' habitat variesfrom Phragmites,Ira, and Smilax to other low shrubs.They did not always nest on the ground. White-faced Ibises nested next to conspecifics,whereas in Glossy Ibises the choice of nearest neighbor was random with respect to the species present. The mean distance to the nearest neighborwas generallyless for GlossyIbis when comparedto White-facedIbis. The spacearound neststhat was devoid of any other nest was examinedgraphically. Nearest neighborswere farther away when they were on the samelevel than when the nearestneighbor was above the nest being examined for both species. The visibility index from the nestsof Glossyand White-faced Ibises was low, and was lessthan the meanvisibility index for heronand egretspecies nesting in the samecolonies. Visibility wasless in the directionof the closestnest comparedto that in other directionsfor all coloniesexamined. In some colonies,the visibility index in the direction of the closestnest was directly related to the distanceto the closestnest. GlossyIbises nested in denserhabitats than did White-faced Ibis, and this was reflected in lower visibility indicesfor GlossyIbis. Possiblereasons for the recent range changesin the two Plegadis speciesare discussed.-- Department of Biology, Livingston College, Rutgers University, New Brunswick, New Jersey 08903. Accepted 15 January 1976. MOST studiesof the breedingbiology, behavior, and ecologyof the family Thres- kiornithidae have only mentioned or describedthe nestinghabitat briefly. We know of no study of the factors determining colony and nest site selectionin any of the ibisesor spoonbills.And, with the exceptionof Jenni's (1969) study on four speciesof heronsduring the breedingseason, little work had beendone on nestingpreferences of heronsand egrets,which often nest with ibisesand spoonbills. We studied the factors determiningcolony and nest site selectionin 2 coloniesof White-faced Ibises (Plegadis chihi) and 5 coloniesof Glossy Ibises (P. falcinellus), with a view to comparingand contrastingcolony and nestsite selectionin these2 closelyrelated species,which may help explain the causesof recentrange changes. Until recently in the Western Hemisphere the Glossy Ibis bred in the Eastern United States and White-faced Ibis bred in the Western United States, Mexico, and southern South America (Palmer 1962, A.O.U. Checklist 1957). Both specieshave been reported to breed in Florida (Brewster 1886) and in Louisiana (Ryder 1967). Historically the White-faced Ibis has had a more extensive,although discontinuous range in the New World (Ryder 1967). Until fairly recently, the breedingrange of GlossyIbis in the United Stateswas limited primarily to Florida (Bull 1974), and the total breedingpopulation was estimatedto be 400 pairs (Palmer 1962).Oberholser (1974) attributed the restrictedrange of GlossyIbis to the fact that its marsh habitat niche was already occupiedby White-faced Ibises. The extensionof the breeding range of GlossyIbis was first noted in the 1940's and 50's (Pearson 1942, Stewart 1957, Stepney and Power 1973). Hailman (1959) summarizedthe breeding expan- 664 The Auk 94: 664-676. October 1977 October1977] White-facedand GlossyIbises 665 sion, which reachednorth to BrigantineNational Wildlife Refugein New Jerseyby 1957. Post et al. (1970) recordedthe first breedingin New York at Jamaica Bay in 1961. By 1973they had increasedto 711 breedingpairs on Long'Island(Buckley and Davis 1973).In 1973Glossy Ibises' breeding range jumped to Maine (Finch 1972,Bull 1974), and in 1975they spreadinto Massachusetts(Finch pers. comm.). Sight records of stragglershave been reportedas far north as St. Pierre off Newfoundland (Finch 1973)and at Bradley'sMarsh near Brandon, Ontario (Goodwinand Bollinger1972). GlossyIbises have alsoundergone a breedingrange expansion into the West Indies and northernSouth America (Gochfeld 1970, 1973). The White-faced Ibis, on the other hand, seemsto be contractingits breeding range, though such range contractionsare poorly documented in the literature. Ryder (1967) summarizedthe breedingcolonies in the United Statesand estimated that in 1965 there were 10,000 breedingpairs. He reporteda declinein breeding numbers in California and attributed it to loss of habitat. Further decline in the numbersbreeding in California was noted by Small (1975). White-faced Ibis are also decliningin Nevada (Zimmerman1975) and Utah (Ryder 1967, Zimmerman1975). A decline in Texas was due to pesticides(Kirke King pers. comm.). Of the Ibis coloniesRyder (1967) lists west of the 89th meridian, 27% were no longer active in 1965. STUDY AREAS AND METHODS We selectedstudy areasto includevaried habitatsin different geographicallocations. Colonies ranged from tules (Scirpus)to small shrubs.In general,colonies were homogeneousphysiognomically. The vegetationeither consistedof all one species(e.g. rulesor Phragmites)or containedseveral species of vegetationintermixed so as to appear similar in structure. Thus the latter type did not contain large patchesof one plant speciessurrounded by tracts of other species.Two heterogeneouscolonies (Danger Island, AransasPass, Texas for the White-facedIbis, and Meadowbrookon LongIsland, New York for the GlossyIbis) were selected for an examinationof generalhabitat preferences, whereas the homogeneous colonieswere selectedto study specificnest site selectionpreferences. Colonies examined were: 1. Lagunade Burgos,Azul, Provinceof BuenosAires, Argentina.This colony,located in a purestand of Scirpussp. (maximumheight 3 m) contained1,500 pairs: 3% Great Egret Casmerodiusalbus, 8% RoseateSpoonbill •tjaia ajaia, 45% SnowyEgret Egretta thula, 18% White-facedIbis, and 26% Brown- hoodedGull Larus maculipennis.Eggs were laid in November. 2. DangerIsland, AransasPass, Texas. The colony,in Borrichiafrutescens,mixed forbs, and Opuntia cactus(maximum height 1.12 m) contained1,500 pairs: 6% GreatBlue Heron•trdea herodias, 2% Reddish Egret Dichromanassarufescens and Little Blue Heron Florida caerulea,37% LouisianaHeron Hyd- ranassatricolor, 30% SnowyEgret, and 25% White-facedIbis. Mixed forbs on DangerIsland included Oenothera drummondii, Ambrosia artemisiifolia, Gaillardia pulchella, Thelespermafilifolium, Machaerantheraphyllocephala, Gavra spp.Helianthus debilis (Jones et al. 1961).Laughing Gulls Larus atricilla nestednearby. Egg-laying extendedfrom February to April. 3. Barrel Island, Brigantine National Wildlife Refuge, New Jersey. This colony, in Iva frutescens (maximum height 1.15 m), contained291 pairs: 12% Great Egret, 13% LouisianaHeron, 36% Snowy Egret,14% Black-crowned Night Heron Nycticorax nycticorax, and 25% Glossy Ibis. Herring Gulls Larus argentatusnested in Spartinaaround the heronry.Egg-laying was in April-May. 4. Meadowbrook in Jones Beach, New York. This colony, in Phragmitescommunis and Myrica pensylvanica(maximum height 3 m) contained90 pairs:64% Black-crownedNight Heronand 36% Glossy Ibis. Herring Gulls nestednearby. Egg-laying was April-May in all the New York colonies. 5. Zach'sBay, JonesBeach, New York. This colony.similar in plant speciesto Meadowbrook,con- tained 500 pairs: 2% LouisianaHeron and Little Blue Heron, 37% Snowy Egret, 7% Black-crowned Night Heron, and 54% GlossyIbis. 6. Loop, JonesBeach, New York. This colony,similar to 4 and 5 in vegetation,contained 139 pairs: 52% Snowy Egret, 7% Black-crownedNight Heron, and 41% GlossyIbis. 7. SeganusThatch, Long Island, New York. This colony,in Smilaxsp. andIva (maximumheight 3.2 m) contained300 pairs:21% Great Egret, 8% Little Blue Heron, 25% SnowyEgret, 19% Black-crowned Night Heron, and 27% GlossyIbis. 666 BURGER AND i•ILLER [Auk, Vol. 94 •oo1 DANG-E R e• WI SE LH GBH AZUL WI SE RS GE G-ULL WHITE-FACED IBIS Fig. 1. Nearestneighbors of White-facedIbises. The brokenbar reflectsthe percent of eachspecies presentand the solidbar the percentnearest neighbor. The diamondreflects a significantdeviation from random.WI = White-facedIbis, SE = SnowyEgret, LH = LouisianaHeron, GBH = GreatBlue Her- on, RS = RoseateSpoonbill, and GE = Great Egret. Fieldworkwas conducted in Argentinain November-December1972 (JB), in Texasin 1974(JB), in New Jerseyin 1973, 1974, and 1975(JB and LMM), and in New York in 1974and 1975 (JB and LMM). Data collectedin all coloniesincluded species of vegetation,height of vegetation,physiognomic charac- teristics,total numberof avian species,and number of individualsbreeding, as well as the relativenumber of eachspecies nesting in the colony.Data collectedon 23 to 65 nestsin eachcolony included: species and heightof vegetationwhere each nest was located, rim heightof the nestabove ground, rim heightfrom the top of the vegetation,nest width and depth,nearest neighbor species, distance to the nearestneighbor, and the distanceto the closestconspecific if a nonconspecificwas the nearestneighbor. The distanceto the nearestneighbor was recordedas the direct linear, horizontal, and vertical distance.We computedthe percentagefrequency of occurrenceas