SEPTEMBER 2011 SHORT COMMUNICATIONS 257

J. Raptor Res. 45(3):257–261 E 2011 The Raptor Research Foundation, Inc.

FOOD HABITS OF THE ROADSIDE HAWK ( MAGNIROSTRIS)DURING THE NONBREEDING SEASON IN THE SOUTHEASTERN PAMPAS OF

ALEJANDRO V. BALADRO´ N1 Laboratorio de Vertebrados, Facultad de Ciencias Exactas y Naturales, Universidad Nacional de Mar del Plata, Funes 3350, B7602AYJ Mar del Plata, Argentina

and

Consejo Nacional de Investigaciones Cientı´ficas y Te´cnicas — CONICET, Rivadavia 1917, C1033AAJ Buenos Aires, Argentina

MARI´A S. BO´ AND ANA I. MALIZIA Laboratorio de Vertebrados, Facultad de Ciencias Exactas y Naturales, Universidad Nacional de Mar del Plata, Funes 3350, B7602AYJ Mar del Plata, Argentina

MARC J. BECHARD Boise State University, Raptor Research Center, Department of Biology, 1910 University Drive, Boise, ID 83706 U.S.A.

KEY WORDS: Roadside Hawk; Buteo magnirostris; Argen- Roadside Hawk delivered mainly small vertebrates tina; diet; nonbreeding diet; pampas; small . (.90%; mainly lizards, frogs, and rodents) to the nest. Studies of raptors’ diet are important for understanding However, Beltzer (1990) analyzed stomach contents of 22 the ecological relationships of the raptors themselves, as Roadside Hawks in Argentina throughout one year, and well as for assessing the influence of these predators on determined that they consumed mostly (77%, community ecology (Marti et al. 2007). Although there is mainly orthopterans). abundant information about food habits of raptor The objectives of our study were to quantitatively de- around the world, data on the trophic ecology of neotrop- scribe the diet of the Roadside Hawk and to estimate its ical raptors are scarce and the diets of many species are food-niche breadth during the nonbreeding season in Ar- poorly known (del Hoyo et al. 1994, Bierregaard 1998, Bo´ gentina. et al. 2007). The Roadside Hawk (Buteo magnirostris) is a widespread METHODS raptor species of Central and , ranging from Our study was conducted in Mar Chiquita Biosphere northern to Rı´o Negro Valley in central Argentina Reserve (southeastern , Argentina; (Thiollay 1994). This medium-sized raptor usually inhabits 37u329 –37u459S; 57u199 –57u269W), a part of the Pampean woodlands and forest edges and, to a lesser extent, open Fitogeographic Region (Cabrera 1971, Viglizzo et al. 2006). fields near woodlands (Canevari et al. 1991). Near the The landscape in this region was dominated by temperate southern boundary of its range, in the pampas of Buenos grasslands, but the original gramineous vegetative commu- Aires Province (Argentina), this raptor is also associated nity has been highly modified by the development of agri- with agroecosystems and grasslands (Narosky and Di Gia- culture (Cabrera 1971, Bilenca and Min˜arro 2004). The como 1993, Mazar-Barnett and Pearman 2001). Reserve comprises a 4600-ha coastal lagoon surrounded Despite being a common species with broad distribu- by native halophytic grasslands, including a diverse array tion, the Roadside Hawk has been little studied and infor- of natural vegetation, such as marshes, coastal dunes, and mation about its food habits is scarce and mostly limited to native forests (Stutz 2001). This habitat heterogeneity sup- qualitative and anecdotal information (e.g., Haverschmidt ports a high faunal diversity, which represents a wide spec- 1962, Massoia 1988). In two studies, the diet of Roadside trum of potential prey for raptors (Martı´nez 2001). Specif- Hawks was quantitatively analyzed. Panasci and Whitacre ically, our sampling was conducted in an area of tala (Celtis (2000) reported breeding-season diet in Guatemala tala) forest at Nahuel Ruca´ Ranch. The tala forest is a type through direct observation of nests, and found that the of thorn forest (Espinal) and is considered threatened due to indiscriminate extraction, cattle grazing, and agriculture 1 Email address: [email protected] (Torres Robles and Tur 2006). This relict patch (6 ha) 258 SHORT COMMUNICATIONS VOL. 45, NO.3 represents the southernmost extreme of Espinal forest in Table 1. Numeric (%F) and biomass (%B) contribution Argentina, and is surrounded by a pond and grazed native of each prey item for the complete sampling period grasslands, which represent potential hunting areas for (nonbreeding seasons 2006–08). Unidentified prey were raptors (Isacch et al. 2000). Adult and juvenile hawks (ap- assigned masses of known prey species of similar size. proximately 10 individuals) were often observed in the tala forest during preliminary surveys. MASS TOTAL During nonbreeding seasons 2006–08 (March to Sep- tember), we collected pellets on 18 occasions at hawks’ PREY ITEM (g) n %F %B roost sites we had previously identified within the tala for- Mammals est. We placed pellets in labeled plastic bags. We later measured the pellets to the nearest mm (length and Rodents width) and soaked them in water to separate osseous and Akodon azarae 28.3 301 62.45 70.34 chitinous materials. We calculated the average number of Calomys spp. 17.4 105 21.78 13.17 prey items per pellet. Oligoryzomys flavescens 26.5 28 5.81 6.86 We classified prey to the lowest possible taxonomic level. Oxymycterus rufus 77.7 4 0.83 1.63 We identified mammals based on skulls and dentaries, Holochilus brasiliensis 266.7 2 0.41 1.21 using identification keys (Bellocq and Kravetz 1983, Go´- Necromys obscurus 40 1 0.21 0.30 mez Villafan˜e et al. 2005) and voucher specimens pre- Unidentified rodents 27.4 17 3.53 4.13 served in the collection of Laboratorio de Vertebrados Marsupials (Facultad de Ciencias Exactas y Naturales, Universidad Na- Monodelphis dimidiata 46.5 16 3.32 2.01 cional de Mar del Plata). Mammals were classified by age (offspring, juvenile, adult or old) based on voucher spec- imens, according to molar wear for cricetine rodents (Bel- Unidentified 34 1 0.21 0.26 locq and Kravetz 1983) and length of dental series for passerines marsupials (Redford and Eisenberg 1992). Amphibians We expressed diet composition as the relative frequency Hypsiboas pulchellus 6 1 0.21 0.05 (number of individuals of each prey item divided by the total number of prey, %F) and as the relative biomass Insects (number of individuals of each prey item multiplied by Orthoptera 1 2 0.41 0.02 the corresponding mean body mass, and divided by the Coleoptera 1 2 0.41 0.02 total biomass consumed, %B). Masses of prey items were Unidentified insects 1 2 0.41 0.02 taken from our unpublished data or from literature (Red- Total 482 ford and Eisenberg 1992, Go´mez Villafan˜e et al. 2005, Vargas et al. 2007). We analyzed the mean size of prey, the food-niche breadth, and the diet similarity among years (Marti et al. 2007). Mean size of prey was estimated as the Geometric RESULTS Mean Weight of Prey (GMWP), which was discriminated by We collected 228 pellets during the nonbreeding sea- age for mammals. Food-niche breadth was calculated using sons for the complete sampling period (75 in 2006, 93 in 2 the Levins’ Index: B 5 1/(Spi ), where pi is the propor- 2007 and 60 in 2008). Pellets were collected at six roost tion of prey item i in the diet, and the Standardized Lev- sites used by about 10 adult and juvenile Roadside Hawks. ins’ Index: Bsta 5 (Bobs 2 Bmin)/(Bmax 2 Bmin), where From these samples, we identified 482 prey, with an aver- Bmin 5 1 and Bmax 5 n; values range from 0 to 1 and allow age of 2.2 6 1.03 prey per pellet (range 5 1–6 prey items). comparisons between samples with different numbers of Pellets averaged 34.1 6 5 mm (range 5 17.4–48.9 mm) in prey items. We examined diet similarity using the Pianka length and 26.7 6 4.1 mm (range 5 14.5–36.8 mm) in 2 2 1/2 Overlap Index: O 5 Spijpik/(Spij pik ) , where pij and width (n 5 185 pellets). pik are the proportions of the prey item i in seasons j and k, The diet of the Roadside Hawk consisted almost exclu- respectively; values range from 0 (no prey item in com- sively of small mammals (98.2%F), especially cricetine ro- mon) to 1 (identical proportional composition; Marti et dents (94.9%F; Table 1). In particular, the field mouse al. 2007). We evaluated statistical differences in prey com- (Akodon azarae) accounted for .55% of the diet for all position among years using the Kendall’s Coefficient of nonbreeding seasons (.63%B). Prey items such as insects Concordance (W) and chi-square test (Zar 1984). Values and amphibians were scarce in the diet of the Roadside of W range from 0 to 1, with higher values indicating Hawk and they were obtained from samples collected at higher similarity. Age composition of mammalian prey in the beginning of the cold season, when these prey items the diet was analyzed by means of G-test (Zar 1984). Values were still available. The majority of small mammals found are expressed as mean 6 SD, followed by range in paren- in hawk’s diet (70.4%F) were species that typically inhabit theses. grasslands (i.e., A. azarae, long-nosed mouse [Oxymycterus SEPTEMBER 2011 SHORT COMMUNICATIONS 259 rufus], dark bolo mouse [Necromys obscurus], and southern Steenhof and Kochert 1985, Jedrzejewski et al. 1994, Figue- short-tailed opposum [Monodelphis dimidiata]), a lower pro- roa Rojas et al. 2003, Sarasola and Negro 2005, Trejo et al. portion (29.1%F) were species that inhabit agroecosystems 2006, Bechard et al. 2010). In our study, the dominance of (i.e., vesper mice [Calomys spp.], pampas rice rat [Oligoryz- small mammals in Roadside Hawk’s diet during the colder omys flavescens]) and only one prey item was a typical wetland nonbreeding season (April–September) coincides with the species (0.004%F; i.e., marsh rat [Holochilus brasiliensis]). period when the abundance of rodents increases due to The Roadside Hawk prey ranged in size from 1 g (in- the recruitment period in the study area (Malizia et al. sects) to 80 g (H. brasiliensis young). Fully 90% of prey 2001). Thus, we do not dismiss the possibility that the represented in the pellets were small-sized rodents, such Roadside Hawk has a more diverse diet during the warmer as A. azarae, O. flavescens, and Calomys spp., all of which breeding season (October-March), when rodent prey weigh ,30 g (Table 1). The age composition of these abundance declines and other prey, including amphibians small rodents was significantly biased toward (a) large in- and invertebrates, are more available. dividuals for A. azarae and O. flavescens (G 5 24.6, df 5 2, P The small species consumed by Roadside Hawks , 0.001 and G 5 8.99, df 5 2, P 5 0.02, respectively) and in our study are typical of the environments which surround (b) juveniles for Calomys spp. (G 5 22.12, df 5 2, P , the tala forest (Malizia et al. 2001). Although this raptor 0.001). In contrast, mammals with mass .30 g, such as seems to hunt in a variety of habitats, it is likely that grass- M. dimidiata, O. rufus, and H. brasiliensis, were uncommon lands and edges are its preferred foraging habitats within (,5%, Table 1) and were represented only by juveniles. As our study area, given the dominance in its diet of A. azarae, a consequence of this dominance of small rodents in the one of the most abundant rodents in grasslands and edges, diet, the GMWP was relatively low (25.2 g 6 0.48 SE). and the presence of M. dimidiata, a species that is almost The food-niche breadth indices were low for the com- endemic to pampean grassland (Redford and Eisenberg plete sampling period (B 5 2.26, n 5 13, and Bsta 5 0.10), 1992, Massoia et al. 2000, Go´mez Villafan˜e 2005). However, reflecting the dominance of a few prey items in the diet of the presence of rodents typically associated with agroecosys- the Roadside Hawk. The Standardized Levins’ Index (Bsta) tems (Calomys spp. and O. flavescens) and wetlands (H. bra- showed little variation among years, varying from 0.15 in siliensis;Go´mez Villafan˜e 2005) suggests some degree of 2006 to 0.17 in 2007 to 0.23 in 2008. opportunism in the hunting habits of this hawk. In addition, no differences in diet composition among We noted that Roadside Hawks consumed mainly juve- years were observed (W 5 0.76; x2 5 27.47, df 5 12, P 5 nile individuals of larger mammals, but preyed on all age 0.007). Dietary overlap was greatest between 2006 and classes of the small-sized mammal species. Although the 2007 (O 5 0.989), intermediate between 2007 and 2008 sample size for large prey was small, our study suggested (O 5 0.973), and least between 2006 and 2008 (O 5 that there may be a maximum prey size Roadside Hawks 0.944). can use, about 80 g, the size of young H. brasiliensis.In addition, the mean prey weight for the Roadside Hawk DISCUSSION (25 g) corresponded with the mean size of the most abun- The Roadside Hawk diet during the nonbreeding season dant cricetine rodents in the study area (Malizia et al. was dominated by small mammals in the southeastern 2001). These findings agreed with those of Panasci and pampas of Argentina. In addition, there were no differenc- Whitacre (2000), who reported that this raptor species es among sampling years, indicating a consistent pattern in preyed mainly on small-sized vertebrate prey. the type of prey taken by this species during the nonbreed- In our study area, this raptor species has experienced a ing season. Our work agreed in part with results reported population increase in recent years, as it was considered by Panasci and Whitacre (2000) during breeding season in rare a few years ago (Martı´nez 2001) and now it seems to Guatemala, where this hawk also consumed many small be a common species (A. Baladro´n unpubl. data). Addi- vertebrates. However, our results differed from those of a tional studies of the Roadside Hawk’s breeding-season diet previous study in Argentina, which included both seasons throughout its entire geographical range are needed to and characterized this species as mainly insectivorous elucidate the latitudinal and seasonal patterns of prey con- (Beltzer 1990). sumption of this predator. Such differences between studies may be due to the different methodological approaches employed, as small HA´ BITOS TRO´ FICOS DE BUTEO MAGNIROSTRIS DU- or soft-bodied prey such as amphibians and insects may RANTE EL PERIODO NO REPRODUCTIVO EN EL SUD- be underrepresented in pellet analyses (Marti et al. ESTE DE LA REGION PAMPEANA DE ARGENTINA 2007). However, such differences may also result from dif- ferences in sampling seasons and areas. The diet of gener- RESUMEN.—Se estudiaron los ha´bitos tro´ficos de Buteo alist raptors usually varies spatially and seasonally, accord- magnirostris durante los periodos no reproductivos entre ing to prey availability (Thiollay 1994). This opportunistic 2006–08 en el sudeste de la regio´n pampeana de Argen- foraging mode has been reported for several , which tina. Colectamos 228 egagro´pilas en seis sitios de descanso generally select the prey that is most locally abundant prey utilizados por adultos y juveniles (aproximadamente 10 or easiest to catch (e.g., Errington and Breckenridge 1938, individuos), e identificamos sus contenidos hasta el nivel 260 SHORT COMMUNICATIONS VOL. 45, NO.3 taxono´mico ma´s bajo posible. B. magnirostris se alimento´ CABRERA, A.L. 1971. Fitogeografı´a de la Repu´blica Argen- principalmente de pequen˜os mamı´feros (98.2%), especial- tina. Boletı´n de la Sociedad Argentina de Bota´nica 19:1–42. mente de roedores (94.9%).´ Items presa tales como aves, CANEVARI, M., P. CANEVARI,R.CARRIZO,G.HARRIS,J.RODRI´- anfibios e insectos fueron encontrados en bajas propor- GUEZ MATA, AND R.J. STRANECK. 1991. Nueva guı´a de las ciones en su dieta. Debido a que el rato´n de campo (Ako- aves . Fundacio´n Acindar, Buenos Aires, Ar- don azarae) fue la presa ma´s importante tanto en nu´mero gentina. (62.5%) como en biomasa (70.3%), (a) la amplitud de DEL HOYO, J., A. ELLIOTT, AND J. SARGATAL. 1994. Handbook nicho tro´fico presento´ valores bajos (I´ndicedeLevins of the birds of the world, Vol 2. New World vultures to Estandarizado 5 0.10) y (b) el taman˜o promedio de presas guineafowl. Lynx Editions, Barcelona, Spain. tambie´n fue relativamente bajo (25.2 g). La composicio´n ERRINGTON, P.L. AND W.J. BRECKENRIDGE. 1938. Food habits de la dieta fue similar para todos los periodos de muestreo of Buteo hawks in north-central United States. Wilson (Coeficiente de Concordancia de Kendall 5 0.76; x2 5 Bulletin 50:113–121. 27.47, gl 5 12, P 5 0.007), mostrando altos valores de FIGUEROA ROJAS, R.A., E.S. CORALES STAPPUNG, AND O.S. solapamiento entre an˜os (I´ndice de Pianka .94%). Nues- ALVARADO. 2003. Diet of the Red-backed Hawk (Buteo tros resultados concordaron con los reportes previos que polyosoma) in a forested area of the Chilean Patagonia indicaban que B. magnirostris consume pequen˜os vertebra- and its relation to the abundance of rodent prey. Hor- dos en Sudame´rica, y evidenciaron tambie´n que la dieta de nero 18:43–52. esta rapaz es ma´s amplia de lo que se habı´a reportado GO´ MEZ VILLAFAN˜ E, I.E., M. MIN˜ O,R.CAVIA,K.HODARA,P. previamente para Argentina, donde era considerada como COURTALO´ N,O.SUA´ REZ, AND M. BUSCH. 2005. Guı´a de una especie principalmente insectı´vora. roedores de la Provincia de Buenos Aires. L.O.L.A. (Lit- [Traduccio´n del equipo editorial] erature of Latin America), Buenos Aires, Argentina. HAVERSCHMIDT, F. 1962. Notes on the feeding habits and ACKNOWLEDGMENTS food of some hawks of Surinam. Condor 64:154–158. We thank P. Urrutia and his family for their courtesy and ISACCH, J.P., M.S. BO´ , AND M.M. MARTI´NEZ. 2000. Food hab- permission to work on his ranch. Our work was supported its of the Striped Owl (Asio clamator) in Buenos Aires by Universidad Nacional de Mar del Plata Fund to Labor- province, Argentina. Journal of Raptor Research 34:235– atorio de Vertebrados (15/E317), and a Conicet Scholar- 237. ship to A. Baladro´n. Equipment for this research was pro- vided through an Idea Wild Grant to A. Baladro´n. We also JEDRZEJEWSKI, W., A. SZYMURA, AND B. 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