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The Condor 95:896-903 0 The Cooper Omlthological Society 1993 SPATIAL AND TEMPORAL ABUNDANCE PATTERNS OF RUDDY QUAIL-DOVES (GEOTRYGON MOKWVi4) NEAR MANAUS, BRAZIL’ PHILIP C. STOUFFER~ AND RICHARD 0. BIERREGAARD, JR. BiodiversityPrograms, NHB 180, SmithsonianInstitution, Washington,DC 20560 Abstract. We analyzed patterns of abundanceof Ruddy Quail-Doves (Geotrygonmon- tana) based on 12 years of mist-net data (457 captures)from terrajirme forest near Manaus in central Amazonian Brazil. Unlike most understory birds at the study site, G. montana varied greatly in abundance. Quail-doves disappeared for months and then reappeared, usually during the wet season.In some months they became one of the most frequently netted birds. Quail-doves avoided isolated forest fragments of one ha, although abundance did not differ among fragments of 10 ha, 100 ha, and continuous forest. Peak abundance varied among years, as did the timing of peak abundance.In general,the annual pattern of quail-dove abundance was correlated with the annual rainfall pattern. Considering all 12 years of data, however, quail-dove abundanceduring a given three-month period was not correlatedwith rainfall during that period, but with rainfall in the sameperiod in the previous year. No quail-doves were recaptured more than a few weeks apart; thus individual birds did not return to the same site from year to year. Since quail-doves feed mainly on fallen fruit, these resultssuggest that they may range over wide areasto exploit regional differences in fruit production. Key words: Amazonian Brazil; Geotrygonmontana; migration; rainfall; Ruddy Quail- Dove:seasonal movements. INTRODUCTION portunity for relatively short distance altitudinal The understoty bird community of terra firme migration suchas is common in Central America forest in central Amazonian Brazil is character- (Levey 1988a, Stiles 1988, Loiselle and Blake ized by permanently resident species that show 199 1) and southeasternBrazil (Sick 1983). A sec- little seasonal variation in abundance as mea- ond reason for the lack of migration may relate sured by mist net captures (Bierregaard 1990a). to the difference in structure of understory com- This area is south of the wintering range for most munities between well-studied sites in Central neotropical migrants (e.g., Blake et al. 1990, Stotz America and Amazonia. Among speciessampled et al. 1992). Austral migrants that winter in equa- by mist-netting, frugivores and nectarivores are torial Brazil include few forest birds (Bierregaard more common in Central America (Karr et al. 1990a). In other tropical regions, forest birds 1990), and are more prone to large- and small- commonly make relatively small scale migra- scalemovements than are insectivores (Terborgh tions to take advantage of seasonalvariation in and Weske 1969, Levey and Stiles 1992). resources,especially differences in flower and fruit The Ruddy Quail-Dove (Geotrygon montana) production as a function of habitat or altitude is one of the few speciesthat exhibit pronounced (Morton 1977, Blake and Loiselle 199 1, Date et variation in abundance in the forests near Ma- al. 199 1, Levey and Stiles 1992). Such move- naus, Brazil, presumably as a result of large-scale ments appear to be much less common in the population movements (Bierregaard 1990a). Amazon Basin than in Central America (Ter- Basedon over seven years of mist net data, Bier- borgh and Weske 1969, Lovejoy 1975, Terborgh regaard (1990a) found it to be more abundant in et al. 1990). the wet season, although there was strong vari- One reason such migrations are less common ation among years. Of the 37 common species in Amazonia may relate to the geographyof the examined, G. montana was the most variable in region. By virtue of its vast expanse with little abundance, disappearing for months at a time. altitudinal variation, Amazonia offers little op In Panama and at La Selva, Costa Rica, abun- dance of G. montana is also highly variable (Stiles and Levey, in press;Karr, pers. observ. cited in I Received12February 1993.Accepted 13 May 1993. 2 Present address: Department of Biological Sci- Karr et al. 1990). ences,Southeastern Louisiana University, Hammond, G. montana is distributed from southern Mex- LA 70401-0814. ico to northern Argentina, including Caribbean P961 RUDDY QUAIL-DOVES IN AMAZONIAN BRAZIL 897 islands (AOU 1983). It is generally found in the The forest is characterized by an average can- interior of moist to wet forests,where it feedson opy height of 30-35 m with occasionalemergents fallen fruits, digestingboth pulp and seeds(Skutch that reach up to 55 m. The understory is rela- 1949; Schubart et al. 1965; Cruz 1974; Davis et tively open and dominated by palms. Density of al. 1985; Hilty and Brown 1986; pers. observ.). fertile understory plants is low compared to other It nests in low vegetation or on decaying stumps neotropical sites, probably becauseof poor soils (Skutch 1949). Active nests were found on the (Gentry and Emmons 1987; seea more complete study site in 1983, 1987, and 1992, and we sus- description ofthe study area in Lovejoy and Bier- pect the bird nested on the site in other years of regaard 1990). Rainfall at Reserva Ducke, 50 km high abundance (D. F. Stotz and A. Whittaker, south of the study site, averagedabout 2.5 m/year pers. comm., pers. observ.). This quail-dove is from 1966 to 1990, with annual peaks from Jan- especially well suited to a study based on mist uary to April (see also RESULTS; rainfall data net captures (see below) as it rarely perches off courtesy of the Departamento de Climatologia the ground and typically flies only one to two m of INPA). above the ground. Results presented here are based on a netting As for many understory birds, preliminary and banding program that began in 1979. All analysis suggestedthat G. montana showsspatial nets were NEBBA-type ATX (36 mm mesh, 12 variation in abundance associatedwith fragment x 2 m). A larger mesh would be more efficient size. Data presentedin Bierregaard and Lovejoy for quail-doves, but since we used the same net (1989; Appendix) suggestthat G. montana de- size at all times, this does not bias our results. creasedin abundance in one ha forest fragments A total of over 55 transects of 100 m (8 nets), in comparison to larger fragments and continu- 200 m (16 nets), or 400 m (30 nets) were netted ous forest. for one day at a time (06:OOto 14:00) at various Here we examine the patterns of spatial and intervals (see Bierregaard 1990a for more de- temporal abundance of G. montana based on 12 tails). Although interspecific differencesin abun- years of mist net capture data. We asked the dance based on mist net capture data must be following questions. How does abundance vary viewed cautiously (see discussion in Karr 198 1, within and among years?Are quail-doves equally Remsen and Parker 1983) we consider the tech- abundant in continuous forest and in patchesof nique effective for determining abundance of one, 10 and 100 ha? How is temporal variation Geotrygon montana based on our knowledge of related to seasonal variation in rainfall? Does the bird’s behavior (see also INTRODUC- temporal variation represent a consistent migra- TION). tion pattern? How can this pattern be explained For most analyses we lumped captures and basedon the ecologicalcharacteristics of the bird? net-hours for three month periods (“quarters” hereafter) that correspondto rainfall seasonality. METHODS Capture rate was distributed bimodally across This study was conducted in terra jirme forest fragment sizesbecause of the low number of cap- 80 km north of Manaus, Brazil (2”3O”S, 6o”W) tures in one ha fragments and frequent absences in the study area of the Biological Dynamics of from larger reserves, so we analyzed fragment Forest Fragments Project. Although some parts size effectsusing a non-parametric test for ran- of the area have been cleared for cattle ranching, domized blocks (Sokal and Rohlf 198 1:446). We providing the opportunity to study the effectsof counted eachquarter with samplesfrom the three fragmentation (e.g., Bierregaard 1990b, Lovejoy fragment sizes and continuous forest as a block, and Bierregaard 1990) these local disturbances thus controlling for variation among quartersand are within an area ofunbroken forestthat extends years. We analyzed relationships among rainfall, for hundreds of km. Here we include data from mass, and abundance using Spearman rank cor- five fragments of one ha, four fragments of 10 relation. Mass data did not violate assumptions ha, two fragments of 100 ha, and continuous forest. Continuous forest data include the frag- of parametric analyses, allowing us to estimate ments before they were isolated by cutting the the variance in mass due to variation among surrounding forest, and five sites that have not years using a model II ANOVA (Sokal and Rohlf been isolated. The greatestdistance between sites 1981:205). We provide details on other tests as is about 60 km. they are used. 898 PHILIP C. STOUFFER AND RICHARD 0. BIERREGAARD, JR. : 0 I\ 1980 1981 1982 1983 1984 1985 1986 1987 ,988 ,989 ,990 I199 YEAR FIGURE I. Monthly capture rate of Geotrygonmomma from October I979 to October 1991, excludingsame- day recaptures.This figure only includes months with >200 net-hours, so gaps in the line represent periods with reduced sampling(e.g., late 1986 and early 1990). RESULTS peaks the abundance of quail-doves relative to other understory birds became quite high; quail- CAPTURE RATE doves were the most frequently netted bird in A total of 457 captures of quail-doves (excluding March 1983 (10.6% of captures) and in March same day recaptures) were recorded in 239,000 1986 (8.3% of captures), and they were the sec- net-hr between October 1979 and October 199 1 ond most commonly netted species in several (Fig.