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J. Merler, M. Diuk-Wasser, R. Quintana (2001)Winter Diet of Dusky-Legged Guan (Penelope obscura) at the Paraná River Delta Region
Studies on Neotropical Fauna and Environment, 36
Iris Ottoni, F. Oliveira, R. Young (2009)Estimating the diet of urban birds: the problems of anthropogenic food and food digestibility.
Applied Animal Behaviour Science, 117
M. Muñoz, G. Londoño, M. Rios, G. Kattan (2007)Diet of the Cauca Guan: Exploitation of a Novel Food Source in Times of ScarcityDieta De Penelope Perspicax: Explotación De Un Recurso Novedoso En Épocas De EscasezCauca Guan Diet
The Condor, 109
J. Altmann (1974)Observational study of behavior: sampling methods.
Behaviour, 49 3
(2008)Terrestrial foot - paddling by a Glaucous - winged Gull
(2006)Diet of the Rustymargined Guan (Penelope superciliaris) in an altitudinal forest fragment of Southeastern Brazil
N Tinbergen (1962)Foot-paddling in gulls
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J. Hoyo, A. Elliott, J. Sargatal, D. Christie (2010)Handbook of the Birds of the World
(1994)Family Numididae (guineafowl), p
S. Mikich (1996)Analise quali-quantitativa do comportamento de Penelope superciliaris (Aves, Cracidae)
Iheringia Serie Zoologia
I. Sazima (2008)Validated cleaner: the cuculid bird Crotophaga ani picks ticks and pecks at sores of capybaras in southeastern Brazil
Biota Neotropica, 8
Ricardo Parrini, Marcos Raposo (2008)Associação entre aves e flores de duas espécies de árvores do gênero Erythrina (Fabaceae) na Mata Atlântica do sudeste do Brasil
Iheringia Serie Zoologia, 98
J del Hoyo, J del Hoyo, A Elliot, J Sargatal (1994)Family Cracidae (chachalacas, guans and curassows)
Handbook of the birds of the world
(1994)Family Cracidae (chachalacas, guans and
I. Sazima, F. Olmos (2009)The Chimango Caracara (Milvago chimango), an additional fisher among Caracarini falcons
Biota Neotropica, 9
K. Frank-Hoeflich, L. Silveira, J. Estudillo-López, A. García-Koch, L. Ongay‐Larios, D. Piñero (2007)Increased taxon and character sampling reveals novel intergeneric relationships in the Cracidae (Aves: Galliformes)
Journal of Zoological Systematics and Evolutionary Research, 45
S B Mikich (2002)A dieta frug�vora de Penelope superciliaris (Cracidae) em remanescentes de foresta estacional semidecidual no centro-oeste do Paran�, Brasil e sua rela��o com Euterpe edulis (Arecaceae)
(2002)A dieta frugívora de Penelope superciliaris
(1985): sampling methods White - breasted guineafowl ( Agelastes meleagrides )
Giulia D'Angelo, I. Sazima (2014)Commensal association of piscivorous birds with foraging otters in southeastern Brazil, and a comparison with such a relationship of piscivorous birds with cormorants
Journal of Natural History, 48
(2015)Uncovering titbits for the ladies : Dusky - legged Guan males stomp out food for their females Ivan Sazima Tinbergen , N . 1962 . Foot - paddling in gulls
M. Muñoz, G. Londoño, M. Rios, G. Kattan (2007)DIET OF THE CAUCA GUAN: EXPLOITATION OF A NOVEL FOOD SOURCE IN TIMES OF SCARCITY
M. Vasconcelos, Camilo Cienfuegos, Lauro Palú (2007)Registros reprodutivos do jacuaçu Penelope obscura Temminck, 1815 (Aves: Cracidae) na porção meridional da Cadeia do Espinhaço, Minas Gerais, Brasil
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S. Cramp (1977)Ostrich to ducks
Revista Brasileira de Ornitologia, 23(3), 315-318 ARTICLE September 2015 Uncovering titbits for the ladies: Dusky-legged Guan males stomp out food for their females 1,2,3 Ivan Sazima Museu de Zoologia, C.P. 6109, Universidade Estadual de Campinas, CEP 13083-970, Campinas, SP, Brazil. Projeto Dacnis, Estrada do Rio Escuro 4754, Sertão das Cotias, CEP 11680-000, Ubatuba, SP, Brazil. Corresponding author: firstname.lastname@example.org Received on 10 December 2014. Accepted on 19 March 2015. ABSTRACT T T: Guans (Cracidae) are vegetarians that feed mostly on fruits but also consume leaves, fl owers, and occasionally invertebrates. Th ey forage mostly by picking food while perched or walking on the ground. I describe here Dusky-legged Guan (Penelope obscura) males that stomp to uncover food in piles of plant litter on the ground. Besides feeding on uncovered titbits, the males feed their mated females that join them on the piles, both by placing food directly in bill or stomping out food that the closely following females pick. Stomping on litter piles to uncover food seems an uncommon and previously unrecorded foraging behaviour. Additionally, uncovering food for following females may be regarded as a type of courtship feeding. KE E EY Y Y-WORDS: Penelope obscura, foraging behaviour, courtship feeding, male-female relationships, Atlantic rainforest, South- eastern Brazil. INTRODUCTION METHODS Guans (Cracidae) are Neotropical galliform birds that Th e Dusky-legged Guan male-female foraging forage singly, in pairs, or small groups, mostly perched interactions were recorded at the edge of the montane on vegetation, although some species or populations Atlantic forest (22°25'54"S, 44°36'55"W, 1.143 m forage on the ground as well (del Hoyo 1994, Sick above sea level) in the Itatiaia range in South-eastern 1997, Zaca et al. 2006, Ottoni et alll. 2009). Guans are Brazil. Th is type of foraging behaviour was observed vegetarians that feed on fruits (their staple diet), leaves, twice in September 2011, and once in September 2013. fl owers, and occasionally invertebrates (del Hoyo 1994, Th roughout the observations, I used the “ad libitum” Merler et alll. 2001, Mikich 2002, Muñoz et alll. 2007, sampling method, which is adequate to record rare events Zaca et alll. 2006, Parrini & Raposo 2008). While (Altmann 1974). Digital photos of the guan foraging foraging on the ground, guans do not rake (scratch) the interactions are housed as vouchers in the Museu de substrate with their feet as several other galliforms do, Zoologia, Universidade Estadual de Campinas (ZUEC). locating the food by sight instead (Sick 1997). However, some cracids occasionally scratch the ground to uncover food (del Hoyo 1994). A recent study indicates that the RESULTS Dusky-legged Guan (Penelope obscura) feeds on human Th e ensuing description is based on the best-documented leftovers in sites close to, or within, urbanised areas (Ottoni et alll. 2009). observation (15 September 2011 at mid-afternoon). A I describe herein the foraging behaviour of Dusky- male Dusky-legged Guan was atop a pile of plant litter legged Guan males that stomp to uncover food hidden stomping on it with the right or the left feet (Figure among piles of plant litter on the ground. Besides feeding 1a). Th is stomping caused the litter to move or slide and uncovered food such as fallen fruits, fresh plant on the uncovered titbits, the males feed their mated females that join them, both by placing food directly in fragments, and insects, which the bird located visually bill or stomping out the food for the closely following and picked with the bill. After about five min of such females. activity by the male, a female approached the pile and Uncovering titbits for the ladies: Dusky-legged Guan males stomp out food for their females Ivan Sazima began to pick food that she located visually, but displayed the female began to follow the stomping male, most of no stomping. Within the first minute of such fora ging, the time partially hidden under his tail (Figure 1d). At the female raised the feathers of her crest and neck, the onset of her following, the male occasionally picked a stretched her dewlap, and closely approached the male food piece and put it in front of the female or on her path (Figure 1b). The male fed her twice, placing food directl y (Figure 1e). Most of the time, however, the male stomped in her bill while she was crouched with neck extended on the pile and the female picked the uncovered food upwards (Figure 1c). This direct feeding occurred shortly right at his feet and under his vent and tail (Figures 1e-f ). after she approached the male and close to the end of Eleven min elapsed from the male stomping on the pile foraging on the litter pile. After the fi rst direct feeding, and his leaving followed by the female (14:43-15:07 h). a b c d FIGURE 1. A Dusky-legged Guan (Penelope obscura) male stomping out food and feeding his mated female. The male lifts his left le g while stomping on a pile of plant litter to uncover food (a); the female (on the left) joins the foraging male – note raised feathers on her head and neck (b); the male (on the left) feeds the female (yellow f ), which is crouched with her neck extended upwards (c); the male continues to stomp on the pile uncovering food and sometimes picking and dropping it in front of the closely following female, partially hidden under his tail (d); the male stomps on the litter and the female picks food close to his feet (e); the male stomps on the edge of the pile, the female still following and picking food (f ). The couple left shortly after. Revista Brasileira de Ornitologia, 23(3), 2015 Uncovering titbits for the ladies: Dusky-legged Guan males stomp out food for their females Ivan Sazima DISCUSSION A A ACKNOWLEDGEMENTS Stomping on litter piles to uncover food seems an I thank Marlies Sazima for help in the field and lovin g uncommon and previously unrecorded foraging behaviour support; Giulia B. D’Angelo for useful comments on the for the Dusky-legged Guan, and other guans as well. first draft; the CNP q for earlier financial su pport. Besides picking food while perched or on the ground, no other type of foraging behaviour seems reported for guans, R R REFERENCES although some cracids occasionally scratch the ground to uncover food (del Hoyo 1994, Sick 1997, Mikich 1996, Altmann, J. 1974. Observational study of behaviour: sampling Zaca et alll. 2006, Ottoni et alll. 2009). Stomping differs methods. Behaviour, 49: 227-267. from scratching (raking), since when the bird stomps, it Arkive 2014a. White-breasted guineafowl ( ( (Agelastes meleagrides s s). touches heavily the substrate with toes widely stretched http://www.arkive.org/white-breasted-guineafowl/agelastes- meleagrides/video-00.html (access on 28 November 2014). and the legs do not move backwards. In contrast, most Arkive 2014b. Red junglefowl (Gallus gallus s s). http://www.arkive.org/ galliforms such as guineafowl and junglefowl scrape red-junglefowl/gallus-gallus/video-08 (access on 28 November away the substrate such as leaf litter and soil to uncover 2014). or unearth food (e.g., Martínez 1994, Arkive 2014a, Burton, R. (1985). Bird behavior. New York, Alfred A. Knopf. b). Stomping seems to be performed only atop a pile of D’Angelo, G. B. & Sazima, I. 2014. Commensal association of piscivorous birds with foraging otters in South-eastern Brazil, plant litter, and thus it would be rarely displayed. Th is and a comparison of such relationship of piscivorous birds with apparently unusual behaviour may be a variation of cormorants. Journal of Natural History, 48: 241-249. scratching displayed by some cracid species (del Hoyo del Hoyo, J. 1994. Family Cracidae (chachalacas, guans and 1994, Sick 1997). Stomping or foot-paddling is part of curassows), p. 310-363. In: del Hoyo, J.; Elliot, A. & Sargatal, J. (eds.). Handbook of the birds of the world. Vol. 2, Ostrich to the foraging repertoire of some gull species and causes ducks. Barcelona, Lynx Edicions. earthworms to surface (Tinbergen 1962, Donnell 2008), Donnell, R. P. 2008. Terrestrial foot-paddling by a Glaucous-winged a function superfi cially similar to that recorded here for Gull. Western Birds, 39: 33-35. the Dusky-legged Guan. Stomping behaviour by this Frank-Hoeflich, K.; S ilveira, L. F.; Estudillo-López, J.; García-Koch, guan may be restricted to few individuals, populations, A. M.; Ongay-Larios, L. & Piñero, D. 2007. Increased taxon and character sampling reveals novel intergeneric relationships in or circumstances, as already commented for the foraging the Cracidae (Aves: Galliformes). Journal of Zoological Systematics tactics of other bird species (e.g., Sazima 2008, Sazima & and Evolutionary Research, 45: 242-254. Olmos 2009, D’Angelo & Sazima 2014). Martínez, I. 1994. Family Numididae (guineafowl), p. 554-567. In: Stomping to uncover food and the associated del Hoyo, J.; Elliot, A. & Sargatal, J. (eds.). Handbook of the birds of the world. Vol. 2, Ostrich to ducks. Barcelona, Lynx Edicions. feeding of a following female appears to be a type of Merler, J. A.; Diuk-Wasser, M. A. & Quintana, R. D. 2001. Winter courtship feeding. In several bird species courtship diet of Dusky-legged Guan (Penelope obscura) at the Paraná River includes the male off ering food to the female (e.g., Burton delta region. Studies on Neotropical Fauna and Environment, 1985, Sick 1997), and Cracinae have a ground display 36: 33-38. courtship, with males feeding females (Frank-Hoeflich et Mikich, S. B. 1996. Análise quali-quantitativa do comportamento de Penelope superciliaris (Aves, Cracidae). Iheringia, Série Zoologia, al. 2007). Most probably, a Dusky-legged Guan female 81: 87-95. approaching a stomping male and leaving with him after Mikich, S. B. 2002. A dieta frugívora de Penelope superciliaris feeding for a while indicates a mated pair. I recorded a (Cracidae) em remanescentes de fl oresta estacional semidecidual copulating pair on 17 October 2007 at the same site I no centro-oeste do Paraná, Brasil e sua relação com Euterpe edulis (Arecaceae). Ararajuba, 10: 207–217. observed the presumed courtship feeding. Additionally, I Muñoz, M.C.; Londoño G. A.; Rios, M. M. & Kattani, G. H. recorded two females with very young chicks on 22 and (2007). Diet of the Cauca Guan; exploitation of a novel food 23 November 2007, which agrees with an account on source in times of scarcity. Th e Condor, 109: 841-851. reproduction of the Dusky-legged Guan in South-eastern Ottoni, I.; Oliveira, F. R. & Y Y Young, R. J. 2009. Estimating the diet Brazil (Vasconcelos et al. 2006). Th us, at the study site of urban birds: Th e problems of anthropogenic food and food digestibility. Applied Animal Behaviour Science, 117: 42–46. breeding of this guan may occur as follows: courtship Parrini, R. & Raposo, M. A. 2008. Associação entre aves e flores and mating in September-October, nesting in October- de duas espécies de árvores do gênero Erythrina (Fabaceae) na November, and rearing chicks in November-December. Mata Atlântica do sudeste do Brasil. Iheringia, Série Zoologia, I suggest here that stomping out food is an unusual 98: 123-128. Sazima, I. 2008. Validated cleaner: the cuculid bird Crotophaga ani type of foraging and when it is displayed in presence of picks ticks and pecks at sores of capybaras in southeastern Brazil. a following female, this type of foraging may function Biota Neotropica, 8: 213-216. as courtship behaviour. Additional studies focused Sazima, I. & Olmos, F. 2009. Th e Chimango Caracara (Milvago on the natural history of the Dusky-legged Guan and chimango), an additional fisher amon g the Caracarini falcons. other cracids (particularly Penelopinae) would test this Biota Neotropica, 9: 403-405. Sick, H. 1997. Ornitologia brasileira. Rio de Janeiro: Editora Nova assumption. Fronteira. Revista Brasileira de Ornitologia, 23(3), 2015 Uncovering titbits for the ladies: Dusky-legged Guan males stomp out food for their females Ivan Sazima Tinbergen, N. 1962. Foot-paddling in gulls. British Birds, Zaca, W.; Silva, W. R. & Pedroni, F. 2006. Diet of the Rusty- 55: 117–120. margined Guan (Penelope superciliaris s s) in an altitudinal forest Vasconcelos, M. F.; Cienfuegos, C. & Palú, L. 2006. Registros fragment of Southeastern Brazil. Ornitologia Neotropical, reprodutivos do jacuaçu Penelope obscura Temminck, 1815 (Aves: 17: 373-382. Cracidae) na porção meridional da Cadeia do Espinhaço, Minas Gerais, Brasil. Lundiana, 7: 145-148. Associate Editor: Cristiano Schetini de Azevedo Revista Brasileira de Ornitologia, 23(3), 2015
Ornithology Research – Springer Journals
Published: Sep 1, 2015
Keywords: Penelope obscura; foraging behaviour; courtship feeding; male-female relationships; Atlantic rainforest; Southeastern Brazil
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