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Ecology and Breeding Habits of the Savanna Hawk in the Llanos of Venezuela

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Ecology and Breeding Habits of the Savanna Hawk in the Llanos of Venezuela Condor 84261-271 0 The Cooper Ornithological Society 1982 ECOLOGY AND BREEDING HABITS OF THE SAVANNA HAWK IN THE LLANOS OF VENEZUELA WILLIAM J. MADER ABSTRACT. -The ecology of the Savanna Hawk (Buteogallusmeridionalis) was studied in the palm-savannas of Venezuela in 1978-1980. Pairs produced only one egg per clutch, no more than one young per calendar year, and had eggsor young in nestsfrom February to December, coinciding largely with the wet season. In comparisonto North American Red-tailed Hawks (Buteojumaicensis), Savanna Hawks had significantly fewer young per pair per year, lower nest success(47.1% vs. 82.5%), mainly due to lower hatching success(64.1% vs. 84.4%) and a lower proportion of pairs laying eggs (73.3% vs. 88.0%). Survival of young Savanna Hawks from hatching to near fledging was 70.4%, and the adult survival rate, 7 1.1%; these values are approximately equal to those for Red-tailed Hawks. The brood size was artificially doubled from one to two at 10 Savanna Hawk nests, but the adults could find only enough food to feed and raise one chick in most cases.Food appeared to be both the ultimate and proximate factor controlling the timing of breeding. The Savanna Hawk has evolved interrelated habits that counter its low fecundity: (1) the capability of renesting after the initial eggs or young have perished; (2) a post-fledging dependency period of four to seven months to help insure survival of the single young; and (3) permanent or long- lived pair-bonds. The ability to renest is enhanced by hunting versatility. Long- lived pair bonds may improve intra-pair activities thereby enhancing the repro- duct&e successof experienced breeders. Hawks and eaglesin the tropics, in contrast to ezuela which examined: (1) a test of the northern latitudes, may have evolved different hypotheses of Lack and Skutch, (2) what reproductive habits in response to greater breeding habits exist to help offset a low repro- diversities of prey and predators, both avian ductive potential inherent with a clutch size of and mammalian, and different climates. Little one, and (3) how the timing and duration of is known, however, about breeding habits and breeding were related to rainfall and food food relationshipsin neotropical falconiforms. resources. Tropical birds generally lay smaller clutches than their temperate counterparts (Lack 1947, STUDY AREA AND METHODS 1948, 1954). Of the several hypotheses that The Savanna Hawk ranges from Panama to have been proposed to explain this contrast, I central Argentina (Blake 1977). I studied a investigated two dealing with the evolution of population for two breeding seasonsand part tropical clutch sizes:Lack ’s (1947) hypothesis ofa third (15 Januarythrough 15 October 1978, of “maximum reproduction” and Skutch’s 25 April through 3 October 1979, and 26 April (1967) of “readjusted reproduction.” The for- through 3 May 1980) on a cattle ranch in Guar- mer theory, which is more widely accepted, ice State, Venezuela. Colleagues continued statesthat birds produce asmany eggsand thus checking nests after I left, from October to offspringas they can feed. The latter maintains December. The habitat was primarily palm that birds produce fewer young than they are savanna, although a few nests were found in capable of feeding becausethe adult survival tree savanna. The region (llanos) has one dry rate is high and the need to recruit breeders is season(generally December through April) and low. Additionally, Skutch believed that a one wet season (May through November), smaller clutch size may, in part, be a result of which have a pronounced effect on the land. heavy predation pressuresin the tropics, such In February for instance, the palm savannas that smaller broods would require fewer feed- were dry and dusty while in August they were ing visits and therefore less risk of nest detec- sometimescovered with over a meter of stand- tion by predators. The question of what selects ing water. for a smaller clutch size in a predatory bird I chose a study area of 9.4 km2 containing has not been addressed. Here, I present the 23 pairs of hawks for which I determined the results of a 16-month study of the Savanna number of eggsand young produced each year. Hawk (Buteogallus meridionalis, following Additional data, including clutch size and Mayr and Cottrell 1979) in the llanos of Ven- nestingsuccess, were gathered on nestslocated 12611 262 WILLIAM J. MADER outside the study area. Nest contents were and exchangesof prey from adult to immature checked periodically using a mirror attached birds. to a pole. I travelled by motorcycle, horse, and on foot. Fifty adult Savanna Hawks were cap- RESULTS tured with balchatri traps (Berger and Mueller 1959). Forty-nine were individually color- PRODUCTIVITY banded, including 13 pairs in which both Savanna Hawks laid eggs from February to members were banded. I could not tell the sex September (n = 116 clutches, Figs. 1 and 2). of adults becausethey differ little in size and Laying peaked in the first part of July 1978 color (mean wing chord = 39.7 cm, range but had no peak in 1979. The number of 31.5-42.3, n = 49 adults; mean weight = 844 clutches in 1979 was nearly equal for each g, range 720-958, y1= 50 adults). Later, I month from April to July, except for a low in determined the sex of several individuals by June. In 1978, hatching occurred from March their behavior. Territories of five neighboring to October with a peak in mid-August. In 1979 pairs(8 of 10 adultswere initially color-banded) nest failures caused two well-defined periods were determined during the dry and wet sea- of hatching, the first in mid-May and the sec- sonsby plotting 50 or more sightingsfor each ond in early September. In both years, I noted pair from February to October 1978. fledging from May to December. Individual Lack’s hypothesis of maximum reproduc- hawks generally did not nest at the same time tion was tested in 1978 by artificially doubling each year. In 1979, marked adults laid eggs, the normal brood size of one, at nine nests on the average, about 30 days from the date during the first week after hatching. The ages they laid in 1978 (Fig. 3). For both years the of the chicks did not differ by more than 1.5 entire breeding season (egg laying-fledging) days. Growth (weight) of two chicks in a spannedabout 250 days (260 days in 1978 and manipulated nest and a chick in a control nest 240 days in 1979) and coincided to a large was measured every five days until fledging. I extent with the wet season(Figs. 1 and 2). doubled the brood size at another nest in 1979 All clutches( yt = 116) had one egg.Nest suc- in order to observe chick interactions and to cess,from egglaying until a chick was at least determine if doubling affected the amount of 5 weeks old, both years combined, was 47.1% prey brought to the nest. In a blind placed level (n = 85 nests, includes 10 renests).The num- with the nest 85 m away, I watched the nest ber of young producedper nest attempt (started for about 44 h from 7 September to 2 October, with a single egg) was 0.47 (n = 85). Of 75 and for 6 h before the secondchick was added. initial attempts in two years, 36 (48.0%) were The average period of observation was 2 h:5 successful;only 4 out of 10 renestsproved suc- min. (range 1:20-2:33). Another blind was cessful (six other renests were noted but not erected at a nest with one chick in 1979 so that checked to determine productivity). For 16 I could collect information about prey (30 h renests, 10 (62.5%) of the initial nestsfailed at observation). Observationsat both blinds were the egg stageand 6 (37.5%) at the chick stage. often interrupted becauseof storms and flood- Of 24 territories thoroughly checked after an ing. I estimated weights of prey from known initial nest failed, 16 (66.7%) pairs renested. weights of comparably sized animals. All No Savanna Hawks attempted to raise two observations were aided by a 32x spotting young in one calendar year. scope. More nests were successful(P < 0.05) in Platforms were built at the top of nine palm 1978 than in 1979. Of 44 initial nestsin 1978, trees (typical nesting sites) containing neststo 25 (56.8%) were successful.Only three ofseven allow a close inspection of chicks and prey (43%) renests were successful. In 1979, 11 remains. Flooding reduced the number of vis- (35.5%) of 3 1 initial nestswere successful.Only its to some nests. Therefore, I considered a one of three renestswas successful.For both nest successfulif a chick was raised to an age years combined, the time of nest failure was of at least five weeks, even though chicks determined for 48 nests,including six renests. fledged(left the nest) at roughly 6.5-7.5 weeks. Thirty-two (66.7%) failed with eggs and 16 I used an incubation period of 39 days (range (33.3%) with young. Hatching success was 37-40, yt = 4) for back-dating some nestsfrom 64.1% (n = 78), while nestling successfrom known hatchingdates to estimatedlaying dates. hatching to at least five weeksof agewas 70.4% Six immature hawks of known fledgingdates (n = 54). were followed to determine the duration of the Nestsbegun during the first half of the breed- post-fledgingdependency period. Dependency ing seasonhad about the same successas nests of a young bird on its parentswas substantiated begun in the last half.
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