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Reproductive behavior of White-tailed Kites (Elanus leucurus) in the Pampas of Argentina

Reproductive behavior of White-tailed Kites (Elanus leucurus) in the Pampas of Argentina Revista Brasileira de Ornitologia 27(3): 153–157. ARTICLE September 2019 Reproductive behavior of White-tailed Kites (Elanus leucurus) in the Pampas of Argentina 1,2 1 1 1 Alejandro V. Baladrón , Matilde Cavalli , Matías G. Pretelli & María S. Bó Laboratorio de Vertebrados, Instituto de Investigaciones Marinas y Costeras (IIMyC), CONICET, Universidad Nacional de Mar del Plata, Mar del Plata, Argentina. Corresponding author: abaladro@mdp.edu.ar Received on 28 March 2019. Accepted on 21 August 2019. ABSTRACT: The White-tailed Kite (E lanus leucurus) is a raptor that has been less studied in the Neotropics than in the Nearctic region, especially in relation to its reproductive behavior. In this study, we report information about the phenology and activity patterns of this raptor at the Pampas of Argentina. We found that White-tailed Kites have a prolonged breeding season, from October to May. Time-activity budgets of mating pairs indicated a pronounced division of roles in parental care between sexes. Females devoted most of their time to nest construction, incubation and chick care (80% of total time) and males to food provisioning and vigilance (70% of total time). We registered 11 cases of prey transfer from the male to the female. In four cases the transfer occurred in flight and in the remaining seven cases while individuals were per ched. Our results agree with general patterns on breeding behavior of White-tailed Kites from North America, suggesting a consistent behavioral pattern throughout the species' distribution. KEY-WORDS: breeding, parental care, raptor, sex roles, time-activity budgets. INTRODUCTION Jaksic 1989, Dunk 1995, Baladrón et al. 2018). The information available about reproductive behavior of the The White-tailed Kite (E lanus leucurus) is a medium- White-tailed Kite is scarce and mostly comes from North sized raptor widely distributed across the Americas America (e.g., Hawbecker 1940 & 1942, Watson 1940). o o (43 N–46 S), from the southeastern United States to the The breeding season covers spring, summer and autumn Patagonia in Argentina and Chile (Figueroa et al. 2006, in its northern range. This prolonged breeding season is Thiollay 2019). This species inhabits open habitats such related to the occurrence of double broods, especially in as wetlands, grasslands and savannas (Fjeldså & Krabbe years of high food abundance (Hawbecker 1940, Dixon 1990, Dunk 1995), but it is also commonly found in et al. 1957, Dunk 1995). There is little information about modified environments such as green areas of cities and parental care, some authors arguing that both parents take villages, agroecosystems, farmlands, and garbage dumps part similarly in the incubation and nest construction (Ferguson-Lees & Christie 2001, Thiollay 2019). Kites activities (Watson 1940, Hawbecker 1942), while others are usually conspicuous during their foraging activities, indicate that females perform most of these tasks (Dixon due to their white coloration and because they search for et al. 1957). The male provides food to the female and prey performing showy flights at medium or low a ltitude chicks throughout the season (Hawbecker 1940, Watson (Mendelsohn & Jaksic 1989, Dunk 1995). 1940, Warner & Rudd 1975). In its southern range, the The feeding ecology is the best known aspect of information available on reproductive behavior is scarce White-tailed Kite's ecology. Most studies performed and, in most cases, anecdotal (Fraga 1984, Jaksic et al. along the species' distribution range agree that this kite 1987), although it is estimated that the breeding season is a specialist predator of small mammals, particularly would also be prolonged in this part of kites' distribution rodents (USA: Hawbecker 1940, Bond 1942, Dixon et al. (de la Peña 2016). 1957, Waian & Stendell 1970, Stendell & Myers 1973; In the framework of a broader study on foraging Chile: Meserve 1977, Schlatter et al. 1980, González- ecology of the White-tailed Kite in the Pampas of Argentina Acuña et al. 2009; Argentina: Leveau et al. 2002, Sarasola (see Baladrón et al. 2018), we registered different activities et al. 2007, 2010, Baladrón et al. 2018; Brazil: Scheibler of kites related to reproductive behavior. In this study, we 2004). This species is characterized as an active-sear ch compiled and analyzed the information collected during predator that uses hovering as its main hunting technique such sampling, emphasizing on reproductive phenology, (Watson 1940, Warner & Rudd 1975, Mendelsohn & parental care, and prey transfer activity. Reproductive behavior of White-tailed Kites Baladrón et al. courtship: interactions between male and female related METHODS to courtship (i.e., vocalizations, material transfer, prey transfer, copulation); 5) in the nest: the individual stays The study was carried out in the southeast of Buenos o o o in the nest; we assumed that the main behavior during Aires province, Argentina (37 32'S–37 55'S; 57 19'W– this activity was incubation or chicks' care (depending 57 30'W), which belongs to the Pampas region (Cabrera on the stage of the breeding cycle). We calculated time- 1971). The landscape in this region is dominated by activity budgets for each sex as the proportion of time grasslands, although the original community of grasses has that each individual devoted to each activity respect to been heavily modified by livestock grazing and agriculture the total time registered each day. Thus, we obtained daily (Soriano et al. 1991, Bilenca & Miñarro 2004). The study budgets for each individual, and then we calculated the area is characterized by its heterogeneity, including a mean values and standard errors of each activity for each diverse array of natural environments, such as grasslands, sex. In addition, we described in a detailed manner the marshes, coastal dunes, and native woodlands, and events of prey transfer from the male to the female and modified environments, such as grazing fields, croplands, quantified t he time that kites employed in prey handling and urban areas (Isacch et al. 2016). and consumption. We identified t he territories of two pairs of kites, from which we obtained data on phenology and behavioral patterns in different years between 2006 and 2008. One RESULTS & DISCUSSION of the groups was located in a woodland area dominated by the native tree known as Tala (Celtis ehrenbergiana) in o o In the southeastern Pampas of Argentina, the White- Nahuel-Rucá ranch (37 37'10''S; 57 25'20''W, hereafter tailed Kite showed a prolonged breeding season. The “tala pair”). The activity of the tala pair was registered starting date of the breeding season was quite variable. during four observation days (26 April 2006: 146 min; 27 We registered copulation events as early as 21 October April 2006: 279 min; 04 May 2006: 256 min; 21 October (tala pair, 2007) and as late as 27 April (periurban pair, 2007: 82 min). The other group was located in a periurban o o 2007). However, we also registered the tala pair making area (Parque Lago village; 37 45'40''S; 57 27'15''W, courtship displays and building the nest as late as 04 hereafter “periurban pair”). The activity of the periurban May 2006. Such delayed starting date would correspond pair was also registered during four observation days to a second nesting attempt. Double-brooding is rare (27 April 2007: 160 min; 22 May 2007: 260 min; 17 in accipitrids, however when the abundance of rodents January 2008: 88 min; 04 April 2008: 312 min). Using is high and prolonged it is known that the White-tailed 10 × 50 binoculars, we recorded the behavior of kites Kite, the Common Black-shouldered Kite (E. axillaris), individually from a fixed position (continuous recor ding and the Black-winged Kite (E. caeruleus) may raise two method; Martin & Bateson 1993, Gaibani & Csermely broods in the same breeding season (Mendelsohn 1983, 2007). From the total sampling time, we quantified only Johnsgard 2009, Thiollay 2019). This behavior has also the reproductive activities of parents in the vicinity of the been recorded for cooperative breeders, such as Harris's nest. We assigned sexes to individuals based mainly on Hawks (Parabuteo unicinctus), for which second brood their behavior, following the behavioral patterns reported attempts will depend on the presence of helpers at the for North America (Watson 1940, Hawbecker 1942, nest (Johnsgard 2009, Thiollay 2019). The final of the Dixon et al. 1957). Although the White-tailed Kite shows breeding season was also variable. We registered fledglings some sexual dimorphism (females are slightly larger and in nest areas as early as 02 February (tala pair, 2007) have darker backs than males; Dunk 1995), these features and as late as 04 April (periurban pair, 2008). Thus, the are of a comparative nature (Hawbecker 1942). Thus, duration of the breeding season included spring, summer we used external characters to differentiate sexes only and autumn, which is in accordance with previous reports when both individuals were seen together. In all cases, from North America (Hawbecker 1940, 1942, Dixon et our assumptions of sex identification (either based on al. 1957). behavior or dimorphism) were positively confirmed Nests were located in the upper part of trees. The during copulation events. All samplings were conducted tala pair placed its nest at ~7.5 m of height in a tala, under good weather conditions (no extreme wind, no which was located at the edge of the woodland (Fig. 1). rain nor fog). This nest was used by kites in previous and later years to We classified t he activities of each individual in one our sampling period (A.V.B. pers. obs.). The periurban of five categories: 1) vigilance: the individual watches pair placed its nest at ~12 m of height in a Eucalyptus spp. over the nest from the air or from a perch; 2) perching: In the same site, we registered another nest (which was the individual remains inactive or performs maintenance abandoned early in the season) at ~4 m in a Pinus spp. activities (cleaning, grooming); 3) nest construction: These results agree with t hose from California, where this the individual carries materials or builds the nest; 4) Revista Brasileira de Ornitologia 27(3): 2019 Reproductive behavior of White-tailed Kites Baladrón et al. kite shows little selectivity in the type and height of trees add new material (Saggese & de Lucca 2001, de la Peña used for nesting (Dixon et al. 1957). 2016), which may be a strategy to save time and energy We registered a total of 550 min of White-tailed at the beginning of the season (Thiollay 2019). Thus, the Kites' reproductive activities. Mating pairs showed a nest building activity that we observe was likely related pronounced division of roles in parental care between to nest “maintenance”, a task that is performed almost sexes, which was revealed by different activity budgets exclusively by the female in most accipitrids, although the between males and females (Fig. 2). From total time male may bring much of the material to her (Rettig 1978, devoted by kites to nest construction, approximately Salvador-Jr. et al. 2008, de Lucca & Saggese 2012). In 98% of time corresponded to females. Previous reports later stages of the breeding cycle, females stood guarding showed mixed results about the participation of sexes in near or into the nest and males devoted most of their nest construction. In California, for example, Watson time to vigilance and prey provisioning (Fig. 2). These (1940) and Hawbecker (1940) reported that both sexes results indicate that incubation and direct feeding of build the nest, but later Dixon et al. (1957) indicated that chicks would be done almost totally by the female. On this task is almost exclusively done by the female, which the other hand, we found that the prey provisioning was is in agreement with our results. However, since both performed in total by the male, which is in agreement pairs nested in previously built nests, in our study we with previous reports for California (Watson 1940). This did not record the complete process of nest construction. pattern of role partitioning between sexes, with the male Many species of accipitrids use old nests, to which they being responsible for most of the hunting and the female Figure 1. Location of one of the nests of White-tailed Kites (Elanus leucurus) in the upper part of a Tala (Celtis ehrenbergiana) in the edge of a Tala woodland (A). Male watching the territory perched near the nest (B) and hunting in a neighboring patch (C). Photo author: M.S.B. (A & B); N. Chiaradia (C). Figure 2. Time-activity budgets of male and female White-tailed Kites (Elanus leucurus) in the Pampas of Argentina. Values are expressed as percentages (mean and standard error). Revista Brasileira de Ornitologia 27(3): 2019 Reproductive behavior of White-tailed Kites Baladrón et al. for most of the parental care in the nest (i.e., incubation, America (Dixon et al. 1957). chicks care, and nest maintenance), is observed in most Studies on raptor behavior are scarce in the accipitrids, except in vultures (Newton 1979, Collopy literature, especially in the Neotropical region. In the case 1984, Pavez 2001, Salvador-Jr. et al. 2008, Thiollay of the White-tailed Kite, there is an asymmetry between 2019). the information available for North and South America We registered 11 events of prey transfer from the (Hawbecker 1940 & 1942, Watson 1940, Warner & male to the female. All events began with the male arriving Rudd 1975, Mendelsohn & Jaksic 1989, Baladrón et al. to the nest area carrying a prey in its talons and emitting 2018). Although based on sporadic records and a modest whistling calls. Watson (1940) reported that the male time of observation, our study show that in the Pampas approaches to the nesting area emitting vocalizations in the White-tailed Kite have a prolonged breeding season order to annunciate his arriving and to coordinate the and show a pronounced division of roles in parental care transfer. Then, the male hovered over the nesting area between sexes. Our findings agree with previous reports holding the prey with his feet downwards. We witnessed for northern populations (Dunk 1995), and suggest four cases of aerial transfer, in which the female leaves her a consistent behavioral pattern throughout species' perch and fluttered a long the male and, when she was distribution. close enough, she positioned behind the male to take the prey with her claws. In the remaining seven cases, the transference occurred in perch. The male arrived carrying ACKNOWLEDGEMENTS the prey and perched next to the female (or the female flies over and per ched next to him), and then she took We thank owners of Nahuel-Rucá ranch for their permit the prey from the side. Watson (1940) reported that both for working in the ranch and to N. Chiaradia for the transfer methods were used in the same frequency by kites photo of the hunting kite. We also thank two anonymous in California. Aerial food exchange seems to be a common reviewers for helpful comments on an earlier version of practice among accipitrids, particularly harriers (Circus) the manuscript. This study was funded by Universidad and kites (Elanus). Although this behavior is clearly linked Nacional de Mar del Plata (EXA 545/11 grant to to sex role division (Watson 1977), the incidence of aerial Laboratorio de Vertebrados) and CONICET (doctoral transfer may respond to differential female requirements scholarship to A.V.B.). The field equipment was provided (e.g., hunger; Dixon et al. 1957), habitat characteristics by Idea Wild (grant to A.V.B.). (e.g., areas of dense vegetal cover; Watson 1977), and predation risk (e.g., for ground nesting species; Negro & REFERENCES Galván 2018). More studies are needed to elucidate the biological significance of this behavior. Baladrón A.V. & Pretelli M.G. 2013. 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Stendell R.C. & Myers P. 1973. White-tailed Kite predation on a Johnsgard P.A. 2009. Birds of the Great Plains: Family Accipitridae fluctuating vole population. Condor 75: 359–360. (hawks, eagles, and harriers). Lincoln: University of Nebraska. Thiollay J.M. 2019. Hawks, eagles (Accipitridae). In: del Hoyo Leveau L.M., Leveau C.M. & Pardiñas U.F.J. 2002. Dieta del Milano J., Elliott A., Sargatal J., Christie D.A. & de Juana E. (eds.). Blanco (Elanus leucurus) en Argentina. Ornitología Neotropical 13: Handbook of the birds of the world alive. Barcelona: Lynx Editions. 307–311. https://www.hbw.com/node/52213 (Access on 05 June 2019). Martin P. & Bateson P. 1993. Measuring behavior: an introductory Waian L.B. & Stendell R.C. 1970. The White-tailed Kite in California guide. Cambridge: Cambridge University Press. with observations of the Santa Barbara population. California Fish Mendelsohn J.M. 1983. Social behavior and dispersion of the Black- & Game 56: 188–198. shouldered Kite. Ostrich 54: 1–18. Warner J.S. & Rudd R.L. 1975. Hunting by the White-tailed Kite Mendelsohn J.M. & Jaksic F.M. 1989. Hunting behaviour of Black- (Elanus leucurus). Condor 76: 226–230. shouldered Kites in the Americas, Europe, Africa and Australia. Watson F.G. 1940. A behavior study of the White-tailed Kite. Condor Ostrich 60: 1–12. 42: 295–304. Meserve P.L. 1977. Food habits of a White-tailed Kite population in Watson A.D. 1977. The Hen Harrier. Berk hamsted: T. & A.D. Poyser. central Chile. Condor 79: 263–265. Negro J.J. & Galván I. 2018. Behavioural ecology of raptors, p. 33– Associate Editor: Cristiano S. Azevedo. Revista Brasileira de Ornitologia 27(3): 2019 http://www.deepdyve.com/assets/images/DeepDyve-Logo-lg.png Ornithology Research Springer Journals

Reproductive behavior of White-tailed Kites (Elanus leucurus) in the Pampas of Argentina

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Revista Brasileira de Ornitologia 27(3): 153–157. ARTICLE September 2019 Reproductive behavior of White-tailed Kites (Elanus leucurus) in the Pampas of Argentina 1,2 1 1 1 Alejandro V. Baladrón , Matilde Cavalli , Matías G. Pretelli & María S. Bó Laboratorio de Vertebrados, Instituto de Investigaciones Marinas y Costeras (IIMyC), CONICET, Universidad Nacional de Mar del Plata, Mar del Plata, Argentina. Corresponding author: abaladro@mdp.edu.ar Received on 28 March 2019. Accepted on 21 August 2019. ABSTRACT: The White-tailed Kite (E lanus leucurus) is a raptor that has been less studied in the Neotropics than in the Nearctic region, especially in relation to its reproductive behavior. In this study, we report information about the phenology and activity patterns of this raptor at the Pampas of Argentina. We found that White-tailed Kites have a prolonged breeding season, from October to May. Time-activity budgets of mating pairs indicated a pronounced division of roles in parental care between sexes. Females devoted most of their time to nest construction, incubation and chick care (80% of total time) and males to food provisioning and vigilance (70% of total time). We registered 11 cases of prey transfer from the male to the female. In four cases the transfer occurred in flight and in the remaining seven cases while individuals were per ched. Our results agree with general patterns on breeding behavior of White-tailed Kites from North America, suggesting a consistent behavioral pattern throughout the species' distribution. KEY-WORDS: breeding, parental care, raptor, sex roles, time-activity budgets. INTRODUCTION Jaksic 1989, Dunk 1995, Baladrón et al. 2018). The information available about reproductive behavior of the The White-tailed Kite (E lanus leucurus) is a medium- White-tailed Kite is scarce and mostly comes from North sized raptor widely distributed across the Americas America (e.g., Hawbecker 1940 & 1942, Watson 1940). o o (43 N–46 S), from the southeastern United States to the The breeding season covers spring, summer and autumn Patagonia in Argentina and Chile (Figueroa et al. 2006, in its northern range. This prolonged breeding season is Thiollay 2019). This species inhabits open habitats such related to the occurrence of double broods, especially in as wetlands, grasslands and savannas (Fjeldså & Krabbe years of high food abundance (Hawbecker 1940, Dixon 1990, Dunk 1995), but it is also commonly found in et al. 1957, Dunk 1995). There is little information about modified environments such as green areas of cities and parental care, some authors arguing that both parents take villages, agroecosystems, farmlands, and garbage dumps part similarly in the incubation and nest construction (Ferguson-Lees & Christie 2001, Thiollay 2019). Kites activities (Watson 1940, Hawbecker 1942), while others are usually conspicuous during their foraging activities, indicate that females perform most of these tasks (Dixon due to their white coloration and because they search for et al. 1957). The male provides food to the female and prey performing showy flights at medium or low a ltitude chicks throughout the season (Hawbecker 1940, Watson (Mendelsohn & Jaksic 1989, Dunk 1995). 1940, Warner & Rudd 1975). In its southern range, the The feeding ecology is the best known aspect of information available on reproductive behavior is scarce White-tailed Kite's ecology. Most studies performed and, in most cases, anecdotal (Fraga 1984, Jaksic et al. along the species' distribution range agree that this kite 1987), although it is estimated that the breeding season is a specialist predator of small mammals, particularly would also be prolonged in this part of kites' distribution rodents (USA: Hawbecker 1940, Bond 1942, Dixon et al. (de la Peña 2016). 1957, Waian & Stendell 1970, Stendell & Myers 1973; In the framework of a broader study on foraging Chile: Meserve 1977, Schlatter et al. 1980, González- ecology of the White-tailed Kite in the Pampas of Argentina Acuña et al. 2009; Argentina: Leveau et al. 2002, Sarasola (see Baladrón et al. 2018), we registered different activities et al. 2007, 2010, Baladrón et al. 2018; Brazil: Scheibler of kites related to reproductive behavior. In this study, we 2004). This species is characterized as an active-sear ch compiled and analyzed the information collected during predator that uses hovering as its main hunting technique such sampling, emphasizing on reproductive phenology, (Watson 1940, Warner & Rudd 1975, Mendelsohn & parental care, and prey transfer activity. Reproductive behavior of White-tailed Kites Baladrón et al. courtship: interactions between male and female related METHODS to courtship (i.e., vocalizations, material transfer, prey transfer, copulation); 5) in the nest: the individual stays The study was carried out in the southeast of Buenos o o o in the nest; we assumed that the main behavior during Aires province, Argentina (37 32'S–37 55'S; 57 19'W– this activity was incubation or chicks' care (depending 57 30'W), which belongs to the Pampas region (Cabrera on the stage of the breeding cycle). We calculated time- 1971). The landscape in this region is dominated by activity budgets for each sex as the proportion of time grasslands, although the original community of grasses has that each individual devoted to each activity respect to been heavily modified by livestock grazing and agriculture the total time registered each day. Thus, we obtained daily (Soriano et al. 1991, Bilenca & Miñarro 2004). The study budgets for each individual, and then we calculated the area is characterized by its heterogeneity, including a mean values and standard errors of each activity for each diverse array of natural environments, such as grasslands, sex. In addition, we described in a detailed manner the marshes, coastal dunes, and native woodlands, and events of prey transfer from the male to the female and modified environments, such as grazing fields, croplands, quantified t he time that kites employed in prey handling and urban areas (Isacch et al. 2016). and consumption. We identified t he territories of two pairs of kites, from which we obtained data on phenology and behavioral patterns in different years between 2006 and 2008. One RESULTS & DISCUSSION of the groups was located in a woodland area dominated by the native tree known as Tala (Celtis ehrenbergiana) in o o In the southeastern Pampas of Argentina, the White- Nahuel-Rucá ranch (37 37'10''S; 57 25'20''W, hereafter tailed Kite showed a prolonged breeding season. The “tala pair”). The activity of the tala pair was registered starting date of the breeding season was quite variable. during four observation days (26 April 2006: 146 min; 27 We registered copulation events as early as 21 October April 2006: 279 min; 04 May 2006: 256 min; 21 October (tala pair, 2007) and as late as 27 April (periurban pair, 2007: 82 min). The other group was located in a periurban o o 2007). However, we also registered the tala pair making area (Parque Lago village; 37 45'40''S; 57 27'15''W, courtship displays and building the nest as late as 04 hereafter “periurban pair”). The activity of the periurban May 2006. Such delayed starting date would correspond pair was also registered during four observation days to a second nesting attempt. Double-brooding is rare (27 April 2007: 160 min; 22 May 2007: 260 min; 17 in accipitrids, however when the abundance of rodents January 2008: 88 min; 04 April 2008: 312 min). Using is high and prolonged it is known that the White-tailed 10 × 50 binoculars, we recorded the behavior of kites Kite, the Common Black-shouldered Kite (E. axillaris), individually from a fixed position (continuous recor ding and the Black-winged Kite (E. caeruleus) may raise two method; Martin & Bateson 1993, Gaibani & Csermely broods in the same breeding season (Mendelsohn 1983, 2007). From the total sampling time, we quantified only Johnsgard 2009, Thiollay 2019). This behavior has also the reproductive activities of parents in the vicinity of the been recorded for cooperative breeders, such as Harris's nest. We assigned sexes to individuals based mainly on Hawks (Parabuteo unicinctus), for which second brood their behavior, following the behavioral patterns reported attempts will depend on the presence of helpers at the for North America (Watson 1940, Hawbecker 1942, nest (Johnsgard 2009, Thiollay 2019). The final of the Dixon et al. 1957). Although the White-tailed Kite shows breeding season was also variable. We registered fledglings some sexual dimorphism (females are slightly larger and in nest areas as early as 02 February (tala pair, 2007) have darker backs than males; Dunk 1995), these features and as late as 04 April (periurban pair, 2008). Thus, the are of a comparative nature (Hawbecker 1942). Thus, duration of the breeding season included spring, summer we used external characters to differentiate sexes only and autumn, which is in accordance with previous reports when both individuals were seen together. In all cases, from North America (Hawbecker 1940, 1942, Dixon et our assumptions of sex identification (either based on al. 1957). behavior or dimorphism) were positively confirmed Nests were located in the upper part of trees. The during copulation events. All samplings were conducted tala pair placed its nest at ~7.5 m of height in a tala, under good weather conditions (no extreme wind, no which was located at the edge of the woodland (Fig. 1). rain nor fog). This nest was used by kites in previous and later years to We classified t he activities of each individual in one our sampling period (A.V.B. pers. obs.). The periurban of five categories: 1) vigilance: the individual watches pair placed its nest at ~12 m of height in a Eucalyptus spp. over the nest from the air or from a perch; 2) perching: In the same site, we registered another nest (which was the individual remains inactive or performs maintenance abandoned early in the season) at ~4 m in a Pinus spp. activities (cleaning, grooming); 3) nest construction: These results agree with t hose from California, where this the individual carries materials or builds the nest; 4) Revista Brasileira de Ornitologia 27(3): 2019 Reproductive behavior of White-tailed Kites Baladrón et al. kite shows little selectivity in the type and height of trees add new material (Saggese & de Lucca 2001, de la Peña used for nesting (Dixon et al. 1957). 2016), which may be a strategy to save time and energy We registered a total of 550 min of White-tailed at the beginning of the season (Thiollay 2019). Thus, the Kites' reproductive activities. Mating pairs showed a nest building activity that we observe was likely related pronounced division of roles in parental care between to nest “maintenance”, a task that is performed almost sexes, which was revealed by different activity budgets exclusively by the female in most accipitrids, although the between males and females (Fig. 2). From total time male may bring much of the material to her (Rettig 1978, devoted by kites to nest construction, approximately Salvador-Jr. et al. 2008, de Lucca & Saggese 2012). In 98% of time corresponded to females. Previous reports later stages of the breeding cycle, females stood guarding showed mixed results about the participation of sexes in near or into the nest and males devoted most of their nest construction. In California, for example, Watson time to vigilance and prey provisioning (Fig. 2). These (1940) and Hawbecker (1940) reported that both sexes results indicate that incubation and direct feeding of build the nest, but later Dixon et al. (1957) indicated that chicks would be done almost totally by the female. On this task is almost exclusively done by the female, which the other hand, we found that the prey provisioning was is in agreement with our results. However, since both performed in total by the male, which is in agreement pairs nested in previously built nests, in our study we with previous reports for California (Watson 1940). This did not record the complete process of nest construction. pattern of role partitioning between sexes, with the male Many species of accipitrids use old nests, to which they being responsible for most of the hunting and the female Figure 1. Location of one of the nests of White-tailed Kites (Elanus leucurus) in the upper part of a Tala (Celtis ehrenbergiana) in the edge of a Tala woodland (A). Male watching the territory perched near the nest (B) and hunting in a neighboring patch (C). Photo author: M.S.B. (A & B); N. Chiaradia (C). Figure 2. Time-activity budgets of male and female White-tailed Kites (Elanus leucurus) in the Pampas of Argentina. Values are expressed as percentages (mean and standard error). Revista Brasileira de Ornitologia 27(3): 2019 Reproductive behavior of White-tailed Kites Baladrón et al. for most of the parental care in the nest (i.e., incubation, America (Dixon et al. 1957). chicks care, and nest maintenance), is observed in most Studies on raptor behavior are scarce in the accipitrids, except in vultures (Newton 1979, Collopy literature, especially in the Neotropical region. In the case 1984, Pavez 2001, Salvador-Jr. et al. 2008, Thiollay of the White-tailed Kite, there is an asymmetry between 2019). the information available for North and South America We registered 11 events of prey transfer from the (Hawbecker 1940 & 1942, Watson 1940, Warner & male to the female. All events began with the male arriving Rudd 1975, Mendelsohn & Jaksic 1989, Baladrón et al. to the nest area carrying a prey in its talons and emitting 2018). Although based on sporadic records and a modest whistling calls. Watson (1940) reported that the male time of observation, our study show that in the Pampas approaches to the nesting area emitting vocalizations in the White-tailed Kite have a prolonged breeding season order to annunciate his arriving and to coordinate the and show a pronounced division of roles in parental care transfer. Then, the male hovered over the nesting area between sexes. Our findings agree with previous reports holding the prey with his feet downwards. We witnessed for northern populations (Dunk 1995), and suggest four cases of aerial transfer, in which the female leaves her a consistent behavioral pattern throughout species' perch and fluttered a long the male and, when she was distribution. close enough, she positioned behind the male to take the prey with her claws. In the remaining seven cases, the transference occurred in perch. The male arrived carrying ACKNOWLEDGEMENTS the prey and perched next to the female (or the female flies over and per ched next to him), and then she took We thank owners of Nahuel-Rucá ranch for their permit the prey from the side. Watson (1940) reported that both for working in the ranch and to N. Chiaradia for the transfer methods were used in the same frequency by kites photo of the hunting kite. We also thank two anonymous in California. Aerial food exchange seems to be a common reviewers for helpful comments on an earlier version of practice among accipitrids, particularly harriers (Circus) the manuscript. This study was funded by Universidad and kites (Elanus). Although this behavior is clearly linked Nacional de Mar del Plata (EXA 545/11 grant to to sex role division (Watson 1977), the incidence of aerial Laboratorio de Vertebrados) and CONICET (doctoral transfer may respond to differential female requirements scholarship to A.V.B.). The field equipment was provided (e.g., hunger; Dixon et al. 1957), habitat characteristics by Idea Wild (grant to A.V.B.). (e.g., areas of dense vegetal cover; Watson 1977), and predation risk (e.g., for ground nesting species; Negro & REFERENCES Galván 2018). More studies are needed to elucidate the biological significance of this behavior. Baladrón A.V. & Pretelli M.G. 2013. 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Journal

Ornithology ResearchSpringer Journals

Published: Sep 1, 2019

Keywords: breeding; parental care; raptor; sex roles; time-activity budgets

References