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ORNITOLOGIA NEOTROPICAL 18: 111–115, 2007 © The Neotropical Ornithological Society

EVIDENCES OF BROOD PARASITISM OF GIANT (MOLOTHRUS ORYZIVORUS) ON SPOT-BREASTED (ICTERUS PECTORALIS) AND STREAK-BACKED (I. PUSTULATUS) ORIOLES

Jeffrey K. McCrary1 & J. Edward Gates2

1College of Natural Resources, Virginia Polytechnic Institute and State University Blacksburg, Virginia 24061, USA. E-mail: [email protected] 2Appalachian Laboratory, University of Maryland Center for Environmental Science, 301 Braddock Road, Frostburg, Maryland 21532, USA.

Parasitismo de cría del Tordo gigante (Molothrus oryzivorus) sobre el Bolsero maculado (Icterus pectoralis) y el Bolsero dorsilistado (I. pustulatus). Key words: Brood parasitism, fitness, , , Molothrus oryzivorus, Spot-breasted Oriole, Icterus pectoralis, Streak-backed Oriole, Icterus pustulatus.

Giant (Molothrus oryzivorus) are obli- Chestnut-headed (P. wagleri), and gate brood parasites specializing on colonial (P. montezuma) (Ori- sock-nesting (Orians 1985, Jaramillo ans 1985, Robinson 1988, Fleischer & Smith & Burke 1999). Two subspecies are recog- 1992, Nava Solorio 1994, Jaramillo & Burke nized, M. o. oryzivora and M. o. impacifica. The 1999). former occurs in and ranges No records to date, however, demonstrate north into and the island of Trinidad; brood parasitism of the Giant Cowbird on the latter ranges from western Panama north any solitary territorial nesting , and to the Carribean coast of and north to they have not been considered among candi- Veracruz (Jaramillo & Burke 1999). Apart date species for this event (Lowther 2006). from a single obscure record of Giant Cow- Here we report on observations made using parasitism of nests of cooperative-breed- field glasses indicating that solitary, territorial- ing Green Jays (Cyanocorax yncas) (Lehmann nesting Spot-breasted Oriole (Icterus pectoralis) 1960), they are known only to place their eggs and Streak-backed Oriole (I. pustulatus) also in the nests of two species of caciques and serve as hosts of the Giant Cowbird. Nesting five species of : Yellow-rumped habitat was broken tropical dry forest with (Cacicus cela), Red-rumped Cacique areas of secondary growth reaching to 15 m (C. haemorrhous), (Psaro- alternating with yards of single-family homes colius decumanus), (P. viridis), in a rural area on the edge of the Chiltepe Russet-backed Oropendola (P. angustifrons), Peninsula Nature Reserve, Nicaragua. This

111 McCRARY & GATES preserve is located on a peninsula that pene- observed on each of the next 4 days for 2–5 h trates Lake Managua, about 20 km north of each day. The area has an incomplete canopy the capital of Managua. Rainfall averages cover in residential yards and secondary under 800 mm yearly, with almost all rainfall growth in nearby fields. The fledgling made concentrated between May and November. begging calls incessantly during all of our The Spot-breasted Oriole inhabits closed observations, and the orioles fed it arthro- forest and areas with sparse tree cover in pods at intervals averaging 4 min. The orioles tropical dry forest from SW Mexico to NW brought arthropods to the fledgling from . It has been documented breeding distances > 80 m. The fledgling remained at from May through July in pendulous nests 8–12 m height in Phyllostylon brasiliensis woven from fine fibrous materials up to 45 (Ulmaceae) and flew occasionally from tree to cm depth, hanging from the fork of a fine tree. A recently built sock-shaped nest, branch 6 to 18 m high (Stiles & Skutch 1989). approximately 40 cm deep at approximately The female builds the nest and incubates the 10 m height, was 60 m from the point at eggs (Orians 1985). The number of eggs is which the cowbird fledgling was first sighted, reported as 3–4 (Jaramillo & Burke 1999). but we could not confirm whether the nest Both sexes feed the nestlings. This species is was used by the orioles feeding the cowbird reported double-brooded in some locales fledgling. There was no evidence of an oriole with the male caring for the first brood while fledgling within 300 m radius of the cowbird the female begins incubating the second. The fledgling. The orioles apparently were no Streak-backed Oriole occupies a similar range longer attending their nest and were not feed- and habitat to the Spot-breasted Oriole, and ing any other fledgling; they appeared to be the two species are often found nesting in dedicated entirely to feeding the single cow- nearby trees. Nesting has been documented bird fledgling. from 3 to 18 m in height in nests from 25 to Beginning on 14 August 2002, we 50 cm in depth, with 3–4 eggs, during May observed a Giant Cowbird fledgling being through June (Stiles & Skutch 1989, Corman fed by two adult Streak-backed Orioles on & Monson 1995). Nest construction is the Chiltepe Peninsula (12°12.868’N, accomplished only by the female and takes 86°18.889’W; 72 m a.s.l.). The vegetation about 25 days. The female alone incubates. structure was similar to that in the prior, The incubation and nestling period appear nearby sighting. As in the prior case, the similar to those of other orioles, taking 12–14 Giant Cowbird fledgling made begging calls days for incubation and about 14 days for ceaselessly and was continuously fed by two fledging (Corman & Monson 1995). Both adult orioles. Residents of the yard next to sexes feed the nestlings. Both the Spot- this sighting claimed that the fledgling had breasted and the Streak-backed orioles have fledged about 1 week earlier from a nest in been reported as possible acceptors of their yard, a pendulous sock about 1 m in (M. aeneus) parasitism length placed approximately 10 m in height, (Sealy et al. 1997, Sealy & Underwood 2004). and had remained in the vicinity, associated On 4 September 1999, we observed both with the adult orioles, during the entire time. male and female adult Spot-breasted Orioles We made another observation at the same site feeding a Giant Cowbird fledgling on the on 19 August 2002, when both adult orioles Chiltepe Peninsula, near Lake Xiloá fed the fledgling for 3 h. On this date, the (12°12.754’N, 86°19.000’W; 59 m a.s.l.). Feed- fledgling occasionally dropped to the ground ing activities by both adult orioles were to eat fruits of Chlorophora tinctoria ().

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Giant Cowbirds have also been reported to 1997). Although the natural range of Monte- eat the fruit of two other moraceous species, zuma Oropendola, the most typical host spe- Ficus trigona and Coussapoa (Robinson 1988). cies in the region, included the Chiltepe We observed the fledgling and orioles again Peninsula, they were not present within at on 20 August for >1 h. As the orioles moved least 10 km of this location (JKM unpubl.), from tree to tree, the cowbird fledgling fol- possibly due to habitat alterations from clear- lowed them while being fed. The orioles ing pasture land and cutting fuelwood. Giant moved >100 m on each foray to retrieve food Cowbirds commonly parasitized Montezuma for the fledgling, and we repeatedly lost con- Oropendola nests in Laguna de Apoyo, some tact with the pair. We did not find any evi- 60 km southeast of Chiltepe Peninsula, where dence of care by the adults for oriole nesting begins at the start of the dry season in fledglings during this period of sightings. December (JKM unpublished data); the para- Skutch (1996) speculated that this common sitism of oriole broods, where nesting occurs observation was likely due to 1) the observer later in the year, would permit more repro- missing the host feeding its own young, 2) ductive cycles by Giant Cowbirds and poten- cowbird fledglings being more conspicuous, tially higher productivity. Giant Cowbirds are or 3) cowbird fledglings quickly outcompeting strong flyers and can easily move between the host young after fledging. We believe that the Chiltepe Peninsula and Laguna de Apoyo, or latter is the best explanation for our observa- other sites, to deposit eggs for breeding in tions. Our last observation of the Giant Cow- alternate seasons. bird fledgling was on 23 August 2002 during The question has been raised whether which time the oriole adults did not appear Giant Cowbird brood parasitism of oropen- after more than 1 h, in spite of repeated call- dolas is parasitism or mutualism (Webster ing by the fledgling, which made locating it 1994). In studies of their parasitism of oro- easy. The fledgling moved from tree to tree pendola nests, elaborate hypotheses of hyper- within a radius of 40 m and on several occa- parasitism and a blurring of the boundary sions, dropped to the ground to feed. We did between parasitism and mutualism (Smith not observe any oriole fledglings in the area 1968, 1983) have been developed regarding (100 m radius) during the entire period of the complex associations between the Giant these observations. Cowbird, its brood host species, wasps and Giant Cowbird fledglings were much bees, and parasitic (Philornis spp.). In larger than the orioles and were distinguish- the smaller, solitary territorial Icterus spp., our able from them and from other cowbirds by observations suggest that parasitism by the their reddish feet, ivory bill, and easily dis- Giant Cowbird may profoundly affect oriole cernable whitish margins on black flight brood success and fitness, given the observed . We considered it very unlikely that concentrated efforts at feeding cowbird fledg- these fledglings were hatched in a known host lings and the absence of host species fledg- nest and were being feed by a species that did lings in both our observations. In addition to not rear the cowbird in its nest, as none of the the Spot-breasted and the Streak-backed ori- previously documented hosts of Giant Cow- oles, the (I. gularis), which is were found in the area (Sealy et al. 1997, slightly larger, also nests in the Chiltepe Pen- Lowther 2006). Furthermore, most auxiliary insula, and two Neotropical migrants, the Bal- feedings of fledgling brood parasites have timore Oriole (I. galbula) and the Orchard been reported in the and the Brown- Oriole (I. spurius), overwinter in this area. The headed Cowbird (M. ater) (Sealy & Lorenzana Altamira Oriole has similar nesting habits to

113 McCRARY & GATES the other two orioles; its nest is similar but geri for their reviews. This is Scientific Con- intermediate in form between the other two tribution Number 4009 of the University of orioles and the oropendolas. The high feeding Maryland Center for Environmental Science, rate of Giant Cowbird fledglings by the ori- Appalachian Laboratory. oles and the lack of oriole fledglings in our observation suggest that Giant Cowbirds may REFERENCES be extremely detrimental to the fitness of nesting oriole species. Corman, T., & G. Monson. 1995. First United It has been noted that conservation States nesting records of the Streak-backed efforts should focus on those species most Oriole. Western Birds 26: 49–53. vulnerable to parasitism, i.e., species that Fleischer, R. C., & N. G. Smith. 1992. Giant Cow- accept parasitism (Sealy & Underwood 2004). bird eggs in the nests of two hosts: the The distribution, ecology, and conservation use of morphology and electrophorectic vari- status of the resident orioles in Nicaragua are ants to identify individuals and species. Condor 94: 572–578. little known, although the Spot-breasted Ori- Gillespie, T. W. 2000. Rarity and conservation of ole has been previously identified as in need forest birds in the tropical dry forest region of of conservation due to its low abundance and Central America. Biol. Conserv. 96: 161–168. limited overall geographic range (Gillespie Jaramillo, A., & P. Burke. 1999. New World black- 2000). Based on our reported evidence of birds: the icterids. Princeton Univ. Press, Princ- brood parasitism of these species by Giant eton, New Jersey. Cowbirds, increased attention to their conser- Lehmann, F. C. 1960. Contribuciones al estudio de vation status may be warranted on the Pacific fauna de . Noved. Colomb. 1: 275– slope of Nicaragua. 276. The evidence we present demonstrates Lowther, P. E. 2006. List of victims and hosts of that Giant Cowbirds parasitize solitary terri- the parasitic cowbirds (Molothrus). The Field Museum, Chicago, Illinois. http://fm1.fieldmu- torial nesting species where colonial sock- seum.org/aa/Files/lowther/CBList.pdf nesting birds are not present, and suggests Nava Solorio, J. 1994. Breeding habits of the Mon- that oriole nests may be parasitized and that tezuma Oropendola (Psarocolius montezuma) at surrogate parental orioles may spend consid- Bacalar, Quintana Roo, Mexico. An. Inst. Biol. erable quantities of energy in chick feeding. Univ. Nac. Autón. Mex. Ser. Zool. 65: 265–274. These results have implications in potential Orians, G. H. 1985. Blackbirds of the Americas. Giant Cowbird habitat and in oriole conser- Univ. of Washington Press, Seattle, Washing- vation. ton. Robinson, S. K. 1988. Foraging ecology and host ACKNOWLEDGMENTS relationships of Giant Cowbird in southeastern Peru. Wilson Bull. 100: 224–235. We wish to thank Kenneth R. McKaye, Sealy, S. G., & J. C. Lorenzana. 1997. Feeding of Lorenzo López, and FUNDECI/GAIA for nestling and fledgling brood parasites by indi- viduals other than the foster parents: a review. field support. Preliminary work on this study Can. J. Zool. 75: 1730–1752. was supported by a Fulbright Fellowship to Sealy, S. G., J. E. Sánchez, R. G. Campos R., & M. JKM. Two anonymous reviewers made valu- Marin. 1997. Bronzed Cowbird hosts: new able comments on an earlier version of this records, trends in host use, and cost of parasit- manuscript. We especially thank Adrianne G. ism. Ornitol. Neotrop. 8: 175–184. Tossas, Adriana Rodríguez-Ferraro, Victoria Sealy, S. G., & T. J. Underwood. 2004. Acceptors D. Paskowski, and Fernando Garbriel Man- and rejectors of cowbird parasitism in the New

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World orioles (Icterus spp.). Ornitol. Neotrop. cago Press, Chicago, Illinois. 15: 331–347. Stiles, F. G., & A. F. Skutch. 1989. A guide to the Skutch, A. F. 1996. Orioles, blackbirds, and their birds of Costa Rica. Comstock Publishing kin: a natural history. Univ. of Arizona Press, Associates, Inc., Cornell Univ. Press, New Tucson, Arizona. York. Smith, N. G. 1968. The advantage of being parasit- Webster, M. S. 1994. Interspecific brood parasitism ized. Nature 219: 690–694. of Montezuma Oropendolas by Giant Cow- Smith, N. G. 1983. Zarhynchus wagleri (Oropendola birds: parasitism or mutualism? Condor 96: cabelicastaña, Oropendola, Chestnut-headed 794–798. Oropendola). Pp. 614–616 in Janzen, D. H. (ed.). Costa Rican natural history. Univ. of Chi- Accepted 10 September 2006.

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