A Comparison of White-Bearded Manakin (Manacus Manacus)Populations and Lek Systems in Suriname and Trinidad
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July1983] ShortCommunications 739 Despite the above arguments, I believe the Red FFRENCH,R. 1973. A guide to the birds of Trinidad Siskin's establishment in Puerto Rico most probably and Tobago. Wynnewood, Pennsylvania, Liv- coincided with the period of its heaviest importation. ingston Publ. Co. This probably would have been in the 1930's when GUNDLACH,J. 1878. Apuntes para la fauna Puerto- the specieswas still relatively common in its native Riquena. Aves. Anales Sociedad Esp. Hist. Nat., lands and there was an extremely high demand for Madrid 2: 141-234, 343-422. it among Canary breeders.Discussions with long-term KING, W. B. 1981. Endangered birds of the world: residents in the region where the Red Siskin occurs The ICBP bird red data book. Washington D.C., could shed additional light on this matter. Smithsonian Institution Press. Regardlessof the length of time that the Red Sis- LEOPOLD, N. F. 1963. Checklist of birds of Puerto kin has inhabited Puerto Rico, its continued survival Rico and the Virgin Islands. Univ. Puerto Rico will depend on protective safeguards. While the Agric. Exp. Sta. Bull. 168. species may have been a will-o'-the-wisp for orni- RAFFAELE,H.A. 1983. A guide to the birds of Puerto thologists, we cannot depend on it remaining so for Rico and the Virgin Islands. San Juan, Puerto pet traders. Rico, Fondo Educativo Interamericano. DE SCHAUENSEE,R.M. 1966. The species of birds of LITERATURE CITED South America and their distribution. Wynne- BIAGGI,V. 1970. Las Aves de Puerto Rico. San Juan, wood, Pennsylvania, Livingston Publ. Co. Puerto Rico, Editorial Universitaria. 1970. A guide to the birds of South Amer- CLAPP,R. B. 1975. Birds imported into the United ica. Wynnewood, Pennsylvania,Livingston Publ. Co. States in 1972. U.S. Fish Wildl. Serv., Spec. Sci. Rept. Wildl. 193. SUNDEVALL,C. J. 1869. Foglarne pa on Portorico, --, & R. C. BANKS. 1973a. Birds imported into efter Hr Hjalmarsons insamlingar framstallda. the United States in 1970. U.S. Fish Wildl. Serv., Ofversigt af Kongl. Vetenskaps-Akademiens Spec. Sci. Rept. Wildl. 164. Forhandlingar. TjugondespetteArgangen 593- 603. --, & ---. 1973b. Birds imported into the WETMORE,A. 1916. Birds of Porto Rico. U.S. Dept. United Statesin 1971. U.S. Fish Wildl. Serv., Spec. Agric. Bull. 326. Sci. Rept. Wildl. 170. DODWELL,G. T. 1976. Encyclopedia of canaries. Received19 August1982, accepted4 January1983. Neptune City, New Jersey,T. F. H. Publications. A Comparison of White-bearded Manakin (Manacus manacus)Populations and Lek Systems in Suriname and Trinidad DEANNA H. OLSON • AND MICHAEL K. McDOWELL 2 •Departmentof Zoologyand 2Schoolof Oceanography,Oregon State University, Corvallis,Oregon 97331-2914 USA The White-bearded Manakin (Manacus manacus)is of Suriname was compared to the Arima Valley pop- a neotropical, primarily frugivorous, lek-mating pas- ulation with respect to population density, morphol- serine. The ecology and behavior of M. m. trinitatis ogy, reproductive behavior, male mating success,and in the Arima Valley, Trinidad has been described lek characteristics. At least two resident males, with (Snow 1962; Lill 1974a, b). Snow (1962) found the display courts near each other, constitute a lek. A Trinidad population density to be unusually high display court is a circular area of forest floor, between with respectto mainland populationsand suggested saplings, cleared of leaf-litter by a male. A resident that this was due to either a high proportion of sec- male successfullydefends his court and a small area ondary forest, causedby limited clearing and lum- surrounding his court against intruding conspecific bering, or reduced interspecific competition on the males. [See Snow (1962) and Lill (1974a) for a more island. Secondaryforest may benefit survival and re- complete description of the lek mating display.] The production, becausemany of the trees abundant in lek characteristicsinvestigated in our study include secondaryforest are prime food sources(Snow 1962) interlek distance, lek area.. intercourt distance, and and there is an abundance of saplings on which the the number of resident males per lek. males depend for their display (Snow 1962, Lill Male dispersion in Suriname and Trinidad was 1974a). compared and related to the lek characteristicsin- There has been little research on M. manacus in vestigated. Male dispersion patterns have been de- mainland South America. In this study, a population scribed as a continuous gradient in the degree of male of M. m. manacusin virgin rain forest in the interior clustering. This gradient ranges from uniform fields 740 ShortCommunications [Auk, Vol. 100 of male territoriesthrough partial clusteringof males Four M. manacusleks were located adjacent to with smaller territories to highly clustered malesand granite plates and domes in dense vegetation char- large intercluster distances,the classicallek disper- acteristic of secondary growth. Males were not ob- sion (Bradbury 1981). We examined male clustering served to display solitarily or in areasdisturbed by and the possible influences of habitat resourcesand treefalls,but the displaysof severalmales were heard population density on male dispersion in M. mana- at two other possible lek locations in dense vegeta- tion near the edge of granite outcrops; these birds The study site was in the Raleigh Falls/Voltzberg were not observed.M. manacusdispersion may be Nature Reserve(approximately 4.5øN, 56.0øW),Suri- limited to areas with vegetation characteristicof sec- name, adjacentto the CoppenameRiver. The primary ondary forest. Secondary growth may supply ade- rain forest was interspersedwith unforested granite quate male display sites,food resources(Snow 1962), outcrops in the form of large, flat plates and domes and nesting sites for manakins. rising to 100 m above the forest canopy Forestchar- The comparisonsof population density, interlek acterized by thick undergrowth and a low canopy distance, lek area, number of males per lek, and in- surrounded the outcrops. This apparent secondary tercourt distance between the two sites are shown in growth may be attributed to rocl•y soil at the edge of Table 1. Snow (1962) determined the population den- the granite outcrops. These areas comprised about sity in the Arima Valley to be 2.7 adults/ha. Lill one-third of our study site, which was located at the (1974b), 10 yr later, estimated that the density had baseof the Voltzberg granite dome within an area of declined by 20% and attributed the decline to the approximately 200 ha. reduction of secondary forest. The Voltzberg adult Data were collected at four leks from December population density was much less than either of these 1980 to May 1981. The averagenearest neighbor, in- Trinidad estimates,confirming Snow's (1962) obser- terlek distancewas determined with adjustmentsfor vation of a lower mainland population density. The topographic relief at the Voltzberg study site and average interlek distancebetween the four Voltzberg comparedwith the average two-dimensional interlek leks was much greater than the average of 11 Arima distance,approximated from Lill (1974b), in the Ari- Valley leks. The four Voltzberg leks consistedof 2, ma Valley. The number of adult resident males per 3, 4, and 5 resident adult males. Lill (1974a) described lek and the location of adult male display courtswere the number of resident males as ranging from 6 to determined by observation during the breeding sea- 50 in a typical Arima Valley lek. Snow (1962) found son. The extent of the breeding seasonin Suriname 205 display courts at seven leks, an average of 29 has not been described. In Trinidad, breeding gen- males per lek. The largest lek in Trinidad had 70 erally occursfrom March to August (Snow 1962, Lill display courts within an area of approximately 18.3 1974a). by 9.2 m (Snow 1962) and was smaller than the av- Snow (1962) estimated the population density of erage polygonal MLA at the Voltzberg site (which adult M. manacusby counting the number of active does not include the two-male lek). MLA was an ex- display courts at the leks to determine the number treme underestimateof the area surrounding the dis- of resident adult males and then calculating the pro- play courts in which resident males actively dis- portion of trapped nonresident males and females to played. The MLA of the three-male lek was 110 m2, trapped resident males. Population density was de- whereas the area in which all territorial and mating termined in the same manner at the Voltzberg study displays occurred was approximately 548 m2. The site from trapping data at leks. nearest-neighbor distance between resident male Males and femalesat two leks were uniquely color display courtsat the four Voltzberg leks ranged from banded, weighed, and measured for right-wing 6.2 to 21.5 m, with an average of 13.6 m, whereas the length. We visited a lek consistingof three color- Arima Valley courtswere, on average, 0.9-4.6 m apart banded resident males daily and observed it from a (Snow 1962). blind on days of high activity. The behaviorsof adult Morphological variation was found between Suri- males, females, and juvenile males at the lek were nam and Trinidad. Adult male wing lengths and recorded during 158 h of observation,between 0600 weights at the Voltzberg site were significantly dif- and 1800.Juvenile maleswere identified by their use ferent (P < 0.001) from those of the Arima Valley of male display patterns.Resident-male territory areas, population (Table 1). determined by locationsof agonisticencounters sur- We observed breeding activity from the onset of rounding individual courts,and lek area in which all our study in mid-December until March. Breeding display activity occurred were determined for this activity was most intense in December and January, three-male lek. The polygonal area enclosedby res- at the start of the short dry season,during which we ident-male display-court centers was used as the logged 114 h of observationtime between 0700 and minimum-lek-area (MLA) estimate for each lek at the 1700.