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Rodriguez and Eberhard 2006

Rodriguez and Eberhard 2006

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Published by marianomx
Breeding observation in Amazon Panamensis
Breeding observation in Amazon Panamensis

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Published by: marianomx on Jan 05, 2009
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The Wilson Journal of Ornithology
118(2):225–236, 2006
ABSTRACT.—We studied the breeding biology of the Panamanian subspecies of the Yellow-crowned Parrot,
 Amazona ochrocephala panamensis
, during 1997–1999 in the province of Chiriquı´, Panama, to provide basicinformation regarding the breeding behavior and reproductive success of these parrots in their natural habitat.We recorded parrot behaviors throughout the reproductive period, monitored nest success, and characterizedoccupied and non-occupied tree cavities. All breeding attempts involved a male-female pair. Clutch size rangedfrom 2 to 4 eggs, which were incubated only by the female, beginning when the first egg was laid. Incubationaveraged 25 days and the eggs hatched asynchronously. During the incubation period, females remained insidethe nest for long periods of time, though they often departed from the nest area during early mornings and lateafternoons, presumably to forage; during this period, males were not observed entering the nest, though theyoften remained nearby. During the nestling period, males contributed significantly to feeding the offspring. Pairsnested in trees that were in good or fair condition, and did not favor cavities in any one tree species. As foundin many other field studies of parrots, breeding success was low. Only 10% (1997–1998) and 14% (1998–1999)of the nests survived poaching and natural predation. Because nest poaching was the primary cause of breedingfailure and poses a serious threat to population viability, we also present data on poaching techniques and thelocal trade of nestling parrots. Overall, the pool of breeding adults is likely made up of aging individuals thatare not being replaced, setting the stage for a rapid population decline.
Received 13 January 2005, accepted 23 November 2005.
The genus
consists of 31 speciesdistributed throughout the Neotropics (Juniperand Parr 1998); however, the breeding biologyof only a few species has been studied (seebelow). Nest poaching and the capture of adult birds for the pet trade, together with hab-itat loss due to deforestation, have contributedto the precipitous decline of 
popu-lations in Central America, South America,and the Caribbean region (Forshaw 1989, Ju-niper and Parr 1998, Wright et al. 2001). Likemany of the eight other subspecies that formthe Yellow-crowned Parrot complex (Juniperand Parr 1998, Eberhard and Bermingham2004),
Amazona ochrocephala panamensis
has not escaped these pressures (Asociacio´nNacional para la Conservacio´n de la Natural-eza 1995, Autoridad Nacional del Ambiente1995a). In Panama, the population of this sub-species has declined considerably due to nest
Escuela de Biologı´a, Univ. Auto´noma de Chiriquı´,David, Panama´.
Smithsonian Tropical Research Inst., Apdo. 2072,Balboa, Panama´.
Current address: Estafeta Universitaria, UNACHI,David, Panama´.
Current address: Biological Sciences Dept. andMuseum of Natural Science, 202 Life Sciences, Lou-isiana State Univ., Baton Rouge, LA 70803, USA.
Corresponding author; e-mail: eberhard@lsu.edu
poaching (Ridgely 1981) and the loss of nest-ing habitat to agricultural and cattle-grazingactivities (Autoridad Nacional del Ambiente1995a, 1995b).The breeding biology of a few
species has been studied in the wild; many of these studies occurred on Caribbean islands(Snyder et al. 1987, Gnam 1991, Rojas-Sua´rez1994, Wilson et al. 1995) while others provideinformation on mainland species (Enkerlin-Hoeflich 1995, Enkerlin-Hoeflich and Hogan1997, Renton and Salinas-Melgoza 1999, andSeixas and Moura˜o 2002). Additional data onbreeding behavior come from studies of cap-tive
A. albifrons
(Skeate 1984) and
A. viridi-genalis
(Wozniak and Lanterman 1984). Over-all, the studies have revealed that females typ-ically spend long periods inside the nest dur-ing the incubation and early nestling periods,and depend, at least to some degree, on beingfed by their mates. Four
species inMexico apparently select nest sites based ontree species, size, cavity height, and entrancesize (Enkerlin-Hoeflich 1995, Renton and Sa-linas-Melgoza 1999).Wright et al. (2001) summarized data frommany field studies and showed that nestpoaching is a principal cause of reproductivefailure in Neotropical parrots, with poachingrates being higher at mainland sites than on
Vol. 118, No. 2, June 2006 
islands, and lower in protected areas (e.g., na-ture reserves). While the impact of nestpoaching on parrot reproductive success isclear, there are few studies that provide infor-mation on specific techniques used by poach-ers.To date, there have been no published stud-ies of the reproductive behavior of 
A. ochro-cephala
in Panama or other parts of its range.Here, we report our observations of the spe-cies’ breeding behavior, describe the charac-teristics of nest sites and nest trees, and quan-tify reproductive success during two breedingseasons. We also present data regarding thepoaching techniques used in the study area.METHODS
Study area.—
Fieldwork was conducted dur-ing the dry season (December–April) of 1997–1998 and 1998–1999 in the lowlands of Corregimiento de San Juan (San Lorenzo dis-trict) of the province of Chiriquı´ in westernPanama. The natural vegetation in the area istropical dry forest (following Holdridge’s[1967] life zone classification) and mangrove,but in many places it has been cleared for ag-riculture and cattle grazing. Annual rainfall is
1,000 mm; mean annual temperature is
C, with mean temperatures of 35
C during the dry and rainy seasons, re-spectively (Instituto de Recursos Hidra´ulicosy Electrificacio´n 1998, 1999). The study areawas located at
N, 82
Wand encompassed an area of 
8,800 ha;
3,875 ha had been partially cleared for ag-riculture and cattle grazing (on haciendas Mir-aflores, El Tekal, and Los Asentamientos deSan Juan), and the remaining 4,925 ha weremangrove. The partially cleared areas stillcontained remnant patches of tropical dry for-est dominated by
Gliricidia sepium
Ery-thrina fusca
trees, the lower-statured
Curatel-la americana
, and palms belonging to the gen-era
(Acosta 1996).
Characterization of nest sites.—
During thefirst breeding season (1997–1998), we onlystudied nests found in the mangrove habitat;in the following season (1998–1999), we ex-tended our nest monitoring to include thosefound in the partially cleared dry forest hab-itat. We found 21 active nests during the 1styear and 42 during the 2nd year. Of the nestsfound in the 2nd year, 14 had been used byparrots during the previous breeding season;therefore, to avoid pseudoreplication, our dataon cavity and nest-tree characteristics repre-sent 49 (and not 63) active nests. In the sec-ond breeding season, 20 of the nests werefound in mangrove habitat, and the remaining22 in the partially cleared dry forest.To find nest cavities, we searched for treeswith cavities, observed parrots flying and vo-calizing in the area, and interviewed local res-idents and field laborers for information aboutnesting parrots. Nests were considered activeif they contained
A. o. panamensis
eggs ornestlings.To determine the availability of cavities, wesearched for additional tree cavities near nesttrees. By searching the area surrounding anoccupied nest tree, we attempted to control forlarger-scale habitat variation (e.g., vegetationdensity, canopy height, distance to feeding ar-eas) that might have influenced cavity choice.All trees within 100 m of each nest tree wereexamined for the presence of large cavities(i.e., cavities similar in size to those occupiedby parrots). For a given nest tree, two of thesurrounding trees found to contain cavitieswere selected at random for inclusion in thesample of unoccupied cavities. If a selectedtree contained more than one cavity, we se-lected one of them at random to provide dataon cavity location and orientation. In the par-tially cleared dry forest habitat, we extendedtwo of these searches beyond 100 m (108 and116 m) in order to find trees with large cavi-ties. Determining that a cavity was similar insize to occupied cavities was admittedly sub- jective; therefore, we do not present any anal-yses comparing the dimensions of occupiedversus unoccupied cavities.We used leaf, flower, and/or fruit samplesto identify the genus and species (where pos-sible) of trees containing cavities. For eachcavity we measured horizontal and verticalwidth of the cavity opening, inside verticaldepth and cavity diameter (measured at thecavity floor), and distance from the ground tothe lower edge of the cavity opening (seeSaunders et al. 1982). Measurements weremade using a 30-m tape to a precision of 0.5cm, and were used to calculate the areas of the cavity entrance and cavity floor. For eachcavity, we noted its location relative to thetree’s structure—branch (cavity completely
 Rodrı´guez and Eberhard 
contained within a branch), trunk (cavity com-pletely contained within the main trunk), andbranch/trunk (cavity at the intersection of abranch and the trunk). We determined the ori-entation of the cavity opening using a com-pass, and measured each tree’s height using aclinometer. We classified the physical condi-tion of each tree—good, fair, poor, or dead—using the scheme outlined by Sauad et al.(1991; see also Saunders et al. 1982).
 Behavioral observations.—
We monitored63 nests during the two breeding seasons: 21during 1997–1998 and 42 during 1998–1999.Of the 63 nests, 5 were selected each year fordetailed behavioral observations of parrots(hereafter referred to as focal nests). In thefirst field season, focal nests were chosen atrandom; during the second field season, nestswere selected on the basis of their accessibil-ity.We made preliminary observations early inthe breeding season (prior to egg-laying) ateach of the focal nests. An observation periodlasted 13 hr (06:00 to 19:00 UTC-5). Eachyear, we watched three of the five focal nestsfor two preliminary observation periods, andthe other two were watched for a single ob-servation period. In most cases (9 of 16 ob-servation periods), we conducted preliminaryobservations prior to capture of the focal in-dividuals.To identify the sex of focal individuals, weused nets (set up at dawn) to capture one orboth members of each focal pair early in thefield season (prior to the onset of breeding).We used nylon (4.5
15 m) and cotton (6
8 m) fishing nets (mist nets were not avail-able) and suspended them using ropes and/orpoles over the nest opening or across a flywayused by the birds. In both years, the sex of each captured individual was identified in thefield by a veterinarian (R. De Obaldı´a) usinga laparoscope. We then marked the female onthe upper chest with Rhodamine B, so that shecould be distinguished from the male in sub-sequent observations. Because the RhodamineB marks faded after several weeks, the birdswere subsequently marked passively by ap-plying dye to the nest opening (see Eberhard1998). This passive marking was done beforethe prior markings had faded completely, sothat the identity of the newly marked birdswas known. With this technique, the birds in-variably marked themselves on different partsof the body with unique patterns, so the maleand female could be distinguished from oneanother.For the remainder of each focal pairsbreeding attempt, we made behavioral obser-vations at
3-day intervals. We observed dur-ing 3-hr periods when the parrots were mostactive (either 06:30–09:30 or 15:45–18:45),following the methodology used in other par-rot studies (e.g., Eberhard 1998, Renton andSalinas-Melgoza 1999). The results reportedhere are based on 859 hr of nest observation(208 hr were preliminary observations). Weobserved nests with the aid of binoculars froma distance of 
15 m (the parrots quickly ha-bituated to the observer’s presence). Duringeach observation period, we noted the follow-ing: time spent by the adults inside the nest;time spent in the nest area (defined as beingin visual range of the observer, which was ap-proximately 50–75 m in the mangrove habitatand approximately 100 m in the partiallycleared dry forest areas); number of other par-rots traveling with the focal individual whenapproaching or departing; and presence of other humans in the nesting area. Other gen-eral observations (allofeeding, allogrooming,vocal and plumage displays, appearances of nestlings at the cavity opening, age at whichyoung left the nest) were noted
ad lib
. Whenadults made short visits to the nest, presum-ably to feed young, we recorded total time inthe nest cavity. Focal nest observations weremade until 6 days after the last chick fledged,or 6 days after a nest was depredated orpoached.Three of the focal nests observed during thefirst breeding season (1997–1998) were incavities that were re-occupied in the followingbreeding season, and were considered focalnests during the 2nd year of the study. Be-cause it is possible that pairs used the samecavity in consecutive years, our data might in-clude some year-to-year pseudoreplication inthe focal-nest behavioral observations. Theadults were not permanently marked, so it wasimpossible to determine whether this oc-curred.For the analysis of behavioral data, we di-vided the breeding season into four stages:pre-laying, laying, incubation, and nestlingperiods. The laying period began with the lay-

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