Facilitation of Sap-Feeding Birds by the White-Fronted Woodpecker in the Monte Desert, Argentina’
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402 SHORT COMMUNICATIONS NELSON,K. S. 1997. Marbled Murrelet (Bruchyram- and enhancementsthrough release 6.11. SAS In- phus marmoratus).In A. Poole and E Gill [eds.], stitute. Inc.. Carv. NC. The birds of North America, No. 276. The Acad- SEALY,S. G. 1972. Adaptive differences in breeding emy of Natural Sciences, Philadelphia, and The biology in the marine bird family Alcidae. Ph.D. American Ornithologists’ Union, Washington, diss., Univ. Michigan, Ann Arbor, MI. DC. SEALY.S. G. 1975a. Asuects of the breeding biologv of the Marbled Muielet in British Columbia. Bi;h PITOCCHELLI,J., J. PIATT, AND M. A. CRONIN.1995. Banding 46:141-154. Morphological and genetic divergence among SEALY,S. G. 1975b. Feeding ecology of the Ancient Alaskan populationsof Brachyrumphusmurrelets. and Marbled Murrelets near Langara Island, Brit- Wilson Bull. 107:235-250. ish Columbia. Can. J. Zool. 53:418-433. RODWAY,M. S. 1990. Status report on the Marbled WHITWORTH,D. L., J. Y. TAKEKAWA,H. CARTER,AND Murrelet in Canada. Committee on the Status of W. R. MCIVER. 1997. A night-lighting technique EndangeredWildlife in Canada, Ottawa, Canada. for at-sea capture of Xantus’ Murrelets. Colonial SAS INSTITUTE.1996. SAS/STAT software: changes Waterbirds20:525-53 1. Thhr Condor 101:402-407 0 The Cooper Ornithological Soc~ty 1999 FACILITATION OF SAP-FEEDING BIRDS BY THE WHITE-FRONTED WOODPECKER IN THE MONTE DESERT, ARGENTINA’ PEDROG. BLENDINGER BiodiversityResearch Group, IADIZA-CRICYT, CC 507, 5500 Mendoza, Argentina, e-mail: [email protected] Abstract. In the Monte desert of South America, fluye de las perforaciones.Otras especiesde aves, de the overall supply of water and food for birds decreas- diferentes grupos ecol6gicos y taxon6micos, aprove- es in the dry, cold season (June through September). than este recurso, que de otro modo raramente esta During this period the White-fronted Woodpecker disponible. La savia es un componenteimportante de (Melanerpes cactorum) drills holes in the trunks and la dieta del Carpintero de 10sCardones y de otras 11 branchesof Prosopisjexuosa and feeds on the exuded especies de aves; durante 10s meses de junio y julio sap. Other bird species,diverse in ecological attributes constituye de1 16% al 83% de las observacionesde and taxonomic affinities, take advantage of this re- forrajeo. La agresi6n por 10s Carpinteros de 10s Car- sourcewhich otherwise would be rarely available. Sap dones reduce significativamenteel tiempo de alimen- is a major constituentof the diet of the White-fronted tacion en las perforaciones con savia de las especies Woodpeckerand 1I other bird species,and sap feeding m&s pequeiias. En la explotacidn de este recurso se comprises between 16% to 83% of foraging observa- revela una compleja combination de interacciones tions made during June and July. Aggression by positivas y negativasentre las aves. Las interacciones White-fronted Woodpeckers significantly reduced the agonisticassugieren que el Carpintero de 10sCardones time smaller bird species spent feeding on sap, indi- compite activamentepor este recurso,mientras que las cating that White-fronted Woodpeckersactively com- otras especies de aves se favorecen al acceder a un pete for this resource. Other bird species profit from recurs0 rice en agua y en azucares. having accessto a resourcerich in water and sugar. Key words: Argentina, competition,facilitation, In Europe and North America, several species of Melanerpes cactorum, Monte desert, sap feeding, woodpeckers feed on the sap flowing from drilled White-frontedWoodpecker. holes (Foster and Tate 1966, MacRoberts and Mac- Roberts 1976, Short 1982). This behavior is perhaps Resumen. En el desiertode1 Monte de Sudamerica, best exemplified by the Yellow-bellied Sapsucker la oferta de agua y aliment0 para las aves disminuye (Sphyrupicusvarius). The sap from holes drilled by durante la estacion seca de inviemo Cjunioa septiem- this speciesattracts, and is used by, other taxa includ- bre). Durante este periodo, el Carpintero de 10s Car- ing insects,birds, and mammals (Foster and Tate 1966, dones (Melanerpes cactorum)taladra 10stroncos y ra- Wiens 1989, Holmes 1990). Use of this food resource mas de Prosopisjexuosa y se alimenta de la savia que may be particularly important to hummingbirds, be- cause they require food of high energy content (Miller and Nero 1983). ’ ’ Received 26 January 1998. Accepted 30 Septem- In South America, two species of woodpeckersof ber 1998. the genus Melanerpes use sap as a food resource:the SHORT COMMUNICATIONS 403 Acorn Woodpecker (M. formicivorus) of the lower of 13 nets were operated periodically, each of them montane wet forest in Colombia (Kattan and Murcia distributed on an area of about 7 ha. 1985, Kattan 1988) and the White-fronted Woodpecker (M. cactorum) (Genise et al. 1993). In the thorn-scrub RESULTS woodland of the semiarid Chaco in Argentina, the During the dry season, White-fronted Woodpeckers White-fronted Woodpecker feeds on sap only during drill holes through the bark and into the phloem of the austral winter; other avian species and insects are Prosopis jenuosu trunks (Fig. 1). Either individually attracted to the drilled holes (Genise et al. 1993). Here or in flocks of up to six birds, White-fronted Wood- I document sap consumption by White-fronted Wood- peckers periodically visited the drillings to feed on the peckers in woodlands occupying the driest portion of exuded sap. Each group of woodpeckers simultaneous- their range, interactions with other bird species that use ly maintained numerous trees with active holes. On a this resource, and seasonal variation in the exploitation few occasions, I observed holes with flowing sap in of sap. If sap is a valuable resource during the dry trunks of Bulnesiu retumu, whereas the two other tree season when both water and food are scarce, then inter- species present in the area, Geqfroea decorticuns and intraspecific agonistic interactions are expected. (chahar) and Ephedru boelckei (patron), were not used by woodpeckers. During the focus sampling period METHODS (June-July), woodpeckers drilled holes only in trunks The study was conducted from November 1996 and branches of P. j?exuosu. I observed only one sap through November 1997 in the Flora and Fauna Re- hole in a trunk of B. retumu in the month of June. serve of Telteca (32”21’S, 68”03’W), Mendoza Prov- However, subsequent observations suggest that, as the ince, Argentina. This reserve is located within the dry season progressed, woodpeckers modified their Monte Phytogeographical Province (Cabrera and Wil- foraging strategy. By early September, they continued link 1980), a narrow arid and semi-arid latitudinal strip to use P. ,jIexuosu, but B. retumu and Lurreu divaricutu in western Argentina, at the foot of the Andes. Telteca shrubs also were frequently drilled, and occasionally is a desert area with marked climatic seasonality, rainy Cupparis atumisqueu. By September, only 33% of the summers, and cold dry winters. Mean annual rainfall observations of sap feeding by all bird species was of is 161 mm (data from Meteorological Program of the P. flexuosa, and 30% each of B. retuma and L. divur- Regional Center for Scientific and Technological In- icatu. vestigations, Mendoza). The landscape is characterized The number of holes in trees and the frequency of by a system of dunes and scrublands dominated by feeding events of the White-fronted Woodpecker Larreu divaricatu Cjarilla), Trichomuriu usillo (usillo), were positively correlated (r3 = 0.65, n = 11, P = and Bulnesiu retamn (retamo). Open woodlands of 0.03). Other bird species clearly took advantage of Prosopis$exuosu (algarrobo dulce) grow in the low- sap flow (Table l), only accessible at recently drilled lying areas between dunes, with a shrub layer domi- woodpecker holes. It is not surprising that the fre- nated by B. retumu, Cuppuris utumisqueu (atamisqui), quency of visits by the other birds also was higher at and Lycium tenuispinosum (Ilaullfn). the most intensely drilled trees (v, = 0.78, n = 11, P In June and July 1997, I made three to nine (mean = 0.005), regardless of the presence or absence of 2 SD = 7.4 ? 2.5) 5-min focal observations of all woodpeckers. birds visiting 11 P. flexuosu trees that were being used Sap consumption by birds was inversely correlated by eight White-fronted Woodpecker flocks. I also re- with rainfall (Fig. 1). During June and July, for bird corded the number of individuals of each species that species using this resource, at least 16% of my for- fed on sap, and the time they spent at the sap tree. aging records was comprised of sap from drilled From November 1996 through October 1997, I made holes (Fig. 2). For each species, I compared the fre- periodic random observations of the foraging behavior quency of feeding at focal sap trees with the species’ of birds showing sap-feeding habits. Whenever possi- relative abundance estimated with mist-nets (Table ble, I collected information on feeding behavior. To 1). Sap consumption was not proportional to bird spe- ensure sample independence, I recorded a single feed- cies abundance in the Telteca Reserve (Fig. 3). At ing event for each individual and, when flocks were least five of these species appeared to actively select involved, only one individual per species (Hejl et al. this resource, including the White-fronted Woodpeck- 1990, Recher and Gebski 1990). I did not knowingly er and Tufted Tit-Spinetail (Leptusthenuru platensis), include the same individual twice in any one day. In whereas other abundant species such as the Common order to diminish overestimation of the most conspic- Diuca-Finch (Diucu diucu) and Picui Ground-Dove uous foraging activities, I only considered the feeding (Columbinu phi) were not observed to use it at all behavior 10 set after a focal bird had been sighted (Fig. 3). (Wiens 1989). White-fronted Woodpeckers defend their feeding The number of sap holes in each tree was estimated holes by driving away smaller bird species attempting by counting the number of trunks and branches, over to use them.