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Abstract
Revista Brasileira de Ornitologia 26(4): 219–226. ARTICLE December 2018 Behavior and seasonal abundance of Neotropic Cormorant Nannopterum brasilianus (Gmelin, 1789) in southeastern, Brazil 1,3 1 2 Tatiane Lima da Silva , Rísia Brígida Gonçalves Cabral & Ildemar Ferreira Programa de Pós-Graduação em Biologia Animal, Instituto de Ciências Biológicas e da Saúde, Universidade Federal Rural do Rio de Janeiro, Rodovia BR 465, km 7, 23897-000, Seropédica, Rio de Janeiro, Brazil. Instituto de Ciências Biológicas e da Saúde, Universidade Federal Rural do Rio de Janeiro, Rodovia BR 465, km 7, 23897-000, Seropédica, Rio de Janeiro, Brazil. Corresponding author: tatianelima@ufrrj.br Received on 18 September 2018. Accepted on 27 November 2018. ABSTRACT: Th e Neotropic Cormorant Nannopterum brasilianus (Gmelin, 1789) is one of the most common and abundant species in Brazil's aquatic environments, but little is known about its natural history. Th is study elaborates and quantitatively evaluates the ethogram and seasonal abundance of this species at Lake Açu, in Rio de Janeiro, Brazil. Between August 2013 and July 2014, behaviors were registered by the ad libitum sampling method and quantifi ed by the continuous recording during 30-min sessions. Th e mean number of sighted individuals per month was 30.41 ± 4.79. Abundance was higher during the dry season (41.5 ± 5.36) compared to the rainy season (19.33 ± 4.81) (t = 3.07, P = 0.01). A total of 42 behaviors were described and grouped into fi ve behavioral categories: maintenance (83.59%), ingestion/excretion (10.67%), locomotion (3.48%), play (1.65%), and agonistic behavior (0.61%). Th e time spent in these categories did not vary signifi cantly between the morning and afternoon periods. Th is study can be used as a starting point for further ethological studies on the species. KEY-WORDS: feeding, Lake Açu, maintenance, Phalacrocoracidae, seasonality. INTRODUCTION near the coast of Rio de Janeiro (Sick 1997, Alves et al. 2011). Th e Phalacrocoracidae family includes about 40 species Although the species is widely distributed and of of long-beaked piscivorous waterbirds, with mostly common occurrence, few studies concerning its natural monochromatic plumage and webbed feet, commonly history have been published. Most include general called Cormorants (Kennedy & Spencer 2014). Despite behavior (e.g., Oliveira & Costa 2003, Sazima 2008, morphological similarities, the typical behaviors among Branco et al. 2009), distribution patterns (e.g., Alves et species in this family may diff er according to the inhabited al. 2011, Silva et al. 2014) and feeding (e.g., Barquete area, due to worldwide distribution (Bernstein & Maxson et al. 2008, Alarcón et al. 2012, Casaux et al. 2012) 1982). Among diff erent behaviors, wing spreading may descriptions. be absent or occur only rarely in some Cormorants, Ethograms are basic tools used in natural history in order to avoid loss of heat in cold regions (Cook & studies (Sakamoto et al. 2009). Th ey consist of a list of Leblanc 2007). behavioral units and their respective descriptions and can Th e Neotropic Cormorant, Nannopterum brasilianus be accompanied by qualitative and quantitative analyses (Gmelin, 1789), the only representative of the family in (Yamamoto & Volpato 2011). Ethograms are reference Brazil, is distributed from the southern United States sources, which may be used in phylogenetic analyses to southern South America (Sigrist 2009). It presents and in the elaboration of hypotheses regarding behavior gregarious habits and inhabits inland waters and seafronts functionality (Kennedy et al. 1996, Gokula 2011). (Sick 1997). Th e species display brown plumage when Behavior descriptions of N. brasilianus are scarce. young and black when adults, with the presence of white Previous studies have been described some aspects related fi loplumes on the sides of the head and neck during the to maintenance (Branco et al. 2009), foraging (Gheler- reproductive period (Bó 1956). In Brazil, they reproduce Costa et al. 2018), agonistic (Oliveira & Costa 2003), in colonies in the Amazon, Pantanal and on islands play (Sazima 2008), feeding associations with piscivorous Revista Brasileira de Ornitologia 26(3): 196–201. ARTICLE September 2018 by Schleidt et al. (1984), Prestes (2000) and Oliveira birds (D'Angelo & Sazima 2013) and dolphins (Santos & Costa (2003). Individuals were classifi ed as young if et al. 2010, Espinoza-Rodríguez et al. 2015), and diving plumage was brown and as adults if plumage was black. behaviors (Quintana et al. 2004). However, an ethogram has not been produced for the species. Adults with white fi loplumes on the sides of the head and Due to the lack and importance of studies related to neck displayed breeding plumage. Th e highest number of individuals observed per month was adopted as a measure natural history, this study elaborates and quantitatively of monthly abundance. analyses Neotropic Cormorant ethogram and seasonal abundance of the species in an artifi cial lake in southeastern Brazil. Ethogram was then compared to others previously Statistical analyses described for Phalacrocoracidae. Data normality (Shapiro-Wilk's test) and homogeneity of variances (Levene's test) were fi rst tested. Th e diff erence between the number of individuals sighted during the METHODS dry (April to September) and rainy (October to March) Study area seasons was analyzed by Student's t-test. Th e behavioral categories were evaluated regarding diff erences in duration between morning and afternoon periods by the Student's Th is study was performed at the campus of the t-test after log-transformation. All tests were performed Universidade Federal Rural do Rio de Janeiro o o (22 46'9.56''S; 43 41'13.12''W) in Seropédica, located using the PAST software v. 3.12 (Hammer et al. 2001), in the Baixada Fluminense district. Th e campus comprises adopting a 5% signifi cance level. Data are presented as 3024 ha (Chiquieri et al. 1995), covering countryside, means ± standard error. grazing areas, Atlantic Forest fragments, spontaneous wetlands, and four lakes, three artifi cial and a natural one RESULTS (Ferreira et al. 2010). Th e climate is classifi ed as “Cw” by Koppen's International System, as humid subtropical with a dry season in winter (Alvares et al. 2013). Th e monthly average of Neotropic Cormorants at Lake Açu was 30.41 ± 4.79 individuals, reaching higher Observations were carried out on an artifi cial lake, and lower values in June 2014 and December 2013, Lake Açu, located near the Agronomy Institute. Th e lake respectively (Fig. 1). Abundances diff ered between seasons comprises an area of 98,538.08 m with a depth of 1.75 m. Th e surrounding vegetation is composed of grasses and (t = 3.07, P = 0.01), higher in the dry season (41.5 ± sparse trees, including Ziziphus joazeiro, Syzygium cumini 5.36) compared to the rainy season (19.33 ± 4.81). Th e population was composed by young individuals, except and S. malaccense; and palms (Arecaceae), including for a record of three adults in reproductive plumage in Caesalpinia echinata, Ficus sp., Mangifera indica, and November 2013. Psidium guajava. Forty-two behaviors were identifi ed and described, Field procedures grouped into fi ve behavioral categories: maintenance (83.59%), ingestion/excretion (10.67%), locomotion Observations were carried out between August 2013 and July 2014. Sixty fi eld trips were performed, totaling 168 h of sampling eff ort, where 100 h comprised the quantitative analyses. Behavioral units were registered by the ad libitum sampling method and quantifi ed by the continuous r ecording (Altmann 1974). Th e observed behaviors were photographed, fi lmed and recorded using spreadsheets. A focal individual was chosen randomly by chance in each session. Th erefore the same individual could or could not be chosen in the next session. During each 30-min session, the duration of each individual behavior was counted in minutes. At the end of each session, 10-min intervals were taken, followed by a new session. A total of 100 sessions were sampled in the morning (06:30 h to 11:59 h) and 100 in the afternoon Figure 1. Abundance (highest monthly value) of Nannopterum (12:00 h to 18:00 h). Th e division and nomenclature brasilianus from August 2013 to July 2014 in the Lake Açu, of each behavior were based on the studies carried out southeastern Brazil. Revista Brasileira de Ornitologia 26(4): 2018 Behavior and abundance of Nannopterum brasilianus Silva et al. (3.48%), play (1.65%) and agonistic behavior (0.61%) Maintenance (Fig. 2). Th ese categories did not vary signifi cantly between the morning and afternoon periods (maintenance: t = Th ese include behaviors related to resting, body care and -0.21, P = 0.83; ingestion/excretion: t = -0.02, P = 0.98; plumage activities (Fig. 3). locomotion: t = -0.85, P = 0.41; play: t = -1.09, P = 0.34; Head scratching: the bird tilts its head to its chest and agonistic behavior: t = -1.17, P = 0.27). and lifts one leg toward it. It then moves the tarsi quickly against the head, in top-to-bottom movements. Th ese movements may also be directed to the chin and throat (Fig. 3A). Back head scratching: the bird moves its neck towards its back, swinging the head against it in quick lateral movements (Fig. 3B). Tarsus scratching: the bird lifts its leg up close to the abdomen and tilts its head towards the tarsi, with rapid half-open beak movements on the tarsi (Fig. 3C). Preening: with its wings half-open and low, the bird moves its half-open beak toward the base of the feathers, aligning them, carrying out this movement for feathers throughout the whole body, tilting its head to the region where the feathers will be aligned (Fig. 3D). Plumage shaking: the bird, with its body leaning Figure 2. Mean duration and standard error of behavioral forward after exiting the water, shakes its whole body in categories during the morning (06:30 h to 11:59 h) and one quick movement, causing excess water to leave the afternoon (12:00 h to 18:00 h) periods (in minutes). Maint - Maintenance, Inges/excre - Ingestion/excretion, Locom - feathers. Th is can be followed or preceded by wing, head Locomotion, Play, and Ag. behav - Agonistic behavior. and tail shaking (Fig. 3E). Figure 3. Behavioral conducts of Nannopterum brasilianus in the maintenance category: (A) Head scratching, (B) Back head scratching, (C) Tarsus scratching, (D) Preening, (E) Plumage shaking, (F) Wing shaking, (G) Wing spreading, (H) Bathing, (I) Sitting resting, (J) Back head resting, (K) Bipedal resting, (L) Unipedal resting, (M) Gular-fl uttering, (N) Neck lowering, (O) Head lowering, (P) Beak opening, (Q) Neck shrinking (adult with breeding plumage), (R) Wing leg stretching. Revista Brasileira de Ornitologia 26(4): 2018 Behavior and abundance of Nannopterum brasilianus Silva et al. Wing shaking: after leaving the water, the bird tilts the bird opens its beak slowly, accompanied by lateral its body forward, opens its wings and shakes them in fast movements of the head (Fig. 3P). and beating movements. Variation: slow wing beating Neck shrinking: the bird shrinks its neck close to (Fig. 3F). the body, forming an “s” (Fig. 3Q). Head shaking: after leaving the water, the bird swings Wing leg stretching: the bird may or may not tilt its head sideways, slow at fi rst, increasing frequency until its body slightly forward, then stretch its wing, followed slowing down again. Th is also occurs during foraging, by the tarsus on the same side of the body (Fig. 3R). after the bird takes its head out of the water. Tail shaking: after leaving the water, the bird leans Ingestion/excretion its body forward and swings its tail horizontally in short and fast movements. Th ese behaviors are related to feeding and hydration Wing spreading: after leaving the water, the bird activities (Fig. 4). opens its wings horizontally and keeps them still (Fig. Foraging: the bird plunges in the water in search 3G). for fi sh by impelling the anterior part of the body, fi rst Bathing: during movement in the water after submerging the head, then the trunk and, fi nally, the foraging, the bird leans its body forward and fl aps its wings tail. Upon returning to the surface, only the neck and against the water, raising its body from the surface. Th is part of the back are exposed. Foraging can be carried out may be followed by diving. Th e bird alternates between individually or in groups, and usually takes place on the displacement in water, dipping, and bathing (Fig. 3H). lakeshore. During group foraging, individuals move in Sitting resting: the bird remains with its body rows, forming a barrier to facilitate fi sh catching (Fig. lowered over its legs, with the abdomen in contact with 4A). the substrate (Fig. 3I). Eating: Th e bird feeds when it emerges after diving Back head resting: the bird tilts its neck towards its during foraging, bringing a fi sh in its beak. It swings its back, positioning the head between the wings with the head sideways to arrange the fi sh into a cranial position in beak tucked beneath scapulars. Eyes remain closed (Fig. the beak, then tilts the head and neck back and swallows 3J). the prey. It may drink water after feeding (Fig. 4B). Bipedal resting: body erect and supported with Drinking: the bird tilts its body under the water, both legs on the substrate, relaxed neck, closed wings and lowers its neck and places its head inside the water (Fig. 4C). lowered tail. Lateral head and gular-fl uttering movements It picks up some water with its beak and vertically raises its may be present (Fig. 3K). head to swallow, with the beak half open (Fig. 4D). Unipedal resting: similar to bipedal resting, but the Defecating: the body is slightly inclined forward, body rests with one leg on the substrate, while the other the feathers close to the cloaca become erect, the tail is is retracted near the abdomen (Fig. 3L). lifted and stool is expelled (Fig. 4E). Gular-fl uttering: the bird remains with its beak half open and vibrates its gular region. Th is behavior is Locomotion prevalent during bipedal resting but can be performed during other behaviors. It is common on hot days (Fig. Th ese are behaviors related to changes in location, 3M). unrelated to feeding (Fig. 5). Neck lowering: the bird lowers its neck vertically Water moving: body in the water with exposed towards the substrate (Fig. 3N). neck, tail and back. Th e bird slowly moves in the water Head lowering: similar to neck lowering, but the by leg movements, without diving (Fig. 5A). bird lowers only its head vertically toward the substrate Open wings water moving: body in a position (Fig. 3O). similar to water moving, but with wings open, high and Beak opening: the bird opens its beak quickly then close to the bird's head as it moves in the water. Th is closes it, as in a yawn. Variation: with its head lowered, behavior precedes fl ight (Fig. 5B). Figure 4. Behavioral conducts of Nannopterum brasilianus in the ingestion/excretion category: (A) Foraging, (B) Eating, (C, D) Drinking, (E) Defecating. Revista Brasileira de Ornitologia 26(4): 2018 Behavior and abundance of Nannopterum brasilianus Silva et al. Figure 5. Behavioral conducts of Nannopterum brasilianus in the locomotion category: (A) Water moving, (B) Open wings water moving, (C) Head dive water moving, (D) Tree branch moving, (E) Falling, (F) Ground moving, (G) Water landing, (H) Water takeoff , (I) Flying. Head dive water moving: body in a position similar tail open. Th e bird performs a beaten fl ight and can glide to water moving, however, the bird repeatedly plunges its for a few seconds (Fig. 5I). head and part of the neck underwater while moving in the water . This can be followed by head shaking when the Play head emerges (Fig. 5C). Tree branch moving: the bird moves laterally on Th ese behaviors are related to the search and movement a branch with its body tilted forward and wings slightly of inanimate objects with the beak (Fig. 6). open and slightly raised, moving one leg at a time in the Single playing: happens on the ground, branches target direction. It can rotate the body 180° to change or water. On the ground, the bird grabs an inanimate position and perform short jumps to move between object (branch, leaf or root) with its beak and swings it nearby branches (Fig. 5D). with its head in lateral movements until the object falls. It Falling: when the bird lands or moves on a branch then picks up the same object and repeats the movement. in trees, it can become unbalanced and fall onto the On branches, the bird moves its head towards branches substrate (the lake, in the case of the present study). To with leaves and tries to pull them out with its beak in try to remain on the branch, it supports its head or body rapid movements. In water, the bird catches an inanimate parts on the branch, with wings and tail open, but this object with its beak and moves in the water with it and does not usually prevent the fall (Fig. 5E). can swing its head laterally (Fig. 6A). Ground moving: the bird moves with one leg at a Group playing: when an individual carries an time in the target direction with its body slightly inclined inanimate object in its beak, another bird approaches, forward, with wings semi-open or closed and raised tail, vocalizes and tries to take the object with its beak (Fig. leaning the body towards the leg that supports it on the 6B). Th is second individual can pick it up from one end ground. Th e bird can rotate its body 180° to change and both then pull on it, like on a tug of war. Other position (Fig. 5F). individuals can vocalize and perform a wing-waving Water landing: body tilted, wings open and raised, display while approaching the bird with the object. In tarsi extended forward and tail open and lowered. Th e bird this display, the bird slightly tilts its body forward, the tail fl ies toward the water, fl aps its wings and slides its legs into is open and raised, and the wings beat slightly . This can be the water surface, with the tarsi being the fi rst part of the accompanied by vocalization (Fig. 6C). body to come into contact with water. When sliding, the Ground searching: the bird tilts its body, lowers its bird slows down, until the abdomen touches the water, neck towards the ground and, with its beak, looks for dry after which the wings close immediately (Fig. 5G). branches or leaves among the vegetation to play with. Th e Water takeoff : with the body in the open wings water bird may walk on the ground or remain in the same place moving position, the bird fl aps its wings and remains with (Fig. 6D). the tail and tarsi in the water. Simultaneously, it moves its legs against the water surface, pushing its body out of the Agonistic behavior water until fl ight takes place (Fig. 5H). Flying: head, neck, trunk, and tail stretched and Th ese are behaviors related to aggressive intra- and aligned horizontally, tarsi extended close to the body and interspecifi c activities (Fig. 6). Revista Brasileira de Ornitologia 26(4): 2018 Behavior and abundance of Nannopterum brasilianus Silva et al. Figure 6. Behavioral conducts of Nannopterum brasilianus in the play and agonistic behavior categories: (A) Single playing, (B) Group playing, (C) Group playing - wing-waving display, (D) Ground searching, (E) Fighting, (F) Ground chasing, (G) Spot stealing, (H) Escaping, (I) Th reat searching. Fighting: body tilted, with wings open or closed. DISCUSSION Th e bird stretches and moves its neck horizontally towards its opponent, and may peck it, preferably on the head or Lake Açu represents a valuable food, rest and overnight neck. Th e gular bag expands and the bird vocalizes (Fig. area for Neotropic Cormorants in urban surroundings. 6E). Th e resident population displayed seasonal variations Ground chasing: the bird pursues the escaping during this monitoring, with greater abundance during opponent in a rapid ground displacement during fi ghting the dry season. A similar pattern was reported for the on the ground. Th e opponent (2) can either face the fi rst nearby Guanabara Bay, southeastern Brazil (Alves et al. individual (1) or continue to run (Fig. 6F). 2011). However, the opposite was recorded in wetlands Spot stealing: when resting on the substrate, in the south and north of Brazil (Branco 2002, Barquete another individual approaches and begins a fi ght. If the et al. 2008, Silva et al. 2014). Th ese variations may be fi rst individual does not change location, it climbs on related to migration between feeding and breeding sites the back of the opponent, fl aps its wings, vocalizes and during the year, which may diff er locally (Sick 1997). pecks the opponent on the neck and head (Fig. 6G). Th e Although adults displaying breeding plumage were attacked bird vocalizes and expands its gular bag. In most found, no reproduction activities were observed. Th e cases, it loses its balance and leaves the area, while the population decreases during the rainy season correspond attacker occupies the free spot. Th e attacking bird may to migration to reproductive sites, usually islands, such as also give up stealing the spot and look for another resting the Tijucas Islands (Alfavaca and Pontuda) in Rio Janeiro place. (Alves et al. 2011). Escaping: displacement of the bird on the ground, Neotropic Cormorant, as well as others cormorants, branch, air or water, away from an intra- or interspecifi c spend most of their daylight time in maintenance activities threat. When escaping by ground, branch, and water, (King et al. 1995, Gwiazda 2000, Branco et al. 2009). Th e the bird moves short distances (1 to 2 m), usually during ethogram raised herein lists behaviors commonly found intraspecifi c agonistic interactions, such as fi ghting or in Phalacrocoracidae family (e.g., Bartholomew et al. spot stealing. Regarding air movement, the bird fl ies at 1968, Oliveira & Costa 2003, Nelson 2005). However, greater distances (> 2 m), preferably during interspecifi c some behaviors seem to have diff erent roles in Neotropic interactions, such as the approach of people or dogs. Cormorant. One of the most common behaviors of When it escapes through the air in areas near water, it the family, wing spreading, is absent in several species propels its body through tarsi movement on the water belonging to the blue-eyed shag complex (Bernstein & surface, similar to water takeoff (Fig. 6H). Maxson 1982), but was frequent in Neotropic Cormorant. Th reat searching: the bird moves its head Th e purpose of such behavior is controversial and may horizontally in the direction of possible threats (Fig. 6I). include thermoregulation (Clark-Jr. 1969), plumage e body can mo Th ve laterally. drying (White et al. 2008) or intraspecifi c display of Vocalization: occurs mainly during agonistic and foraging success (Jones 1978). At Lake Açu, this behavior play interactions. Th e bird opens its beak, expands its was carried out with wet plumage, followed or not by gular bag and vocalizes repeatedly with the proximity of feeding and associated with behaviors to remove excess another individual (oak, oak, oak ...). Variation: small water, indicating a primary plumage drying purpose. hoarse (oak) during resting or foraging. Behaviors associated with reproduction in other Revista Brasileira de Ornitologia 26(4): 2018 Behavior and abundance of Nannopterum brasilianus Silva et al. Cormorant Phalacrocorax capensis ‒ Part 1. Madoqua 9: 5‒30. cormorants were recorded in non-reproductive situations Bó N.A. 1956. Observaciones morfológicas y etológicas sobre el among young individuals, indicating possible alternative Biguá. Hornero 10: 147–157. purposes. Th e wing-waving display performed during Branco J.O. 2002. Flutuações sazonais na abundância de Phalacrocorax group playing is a male advertising carried out during brasilianus (Gmelin) no estuário do Saco da Fazenda, Itajaí, Santa courtship to attract females or claim possession of the nest Catarina, Brasil. Revista Brasileira de Zoologia 19: 1057‒1062. Branco J.O., Evangelista C.L., Lunardon-Branco M.J., Azevedo- (van Tets 1965, Childress & Bennun 2002). However, it can Júnior S.M. & Larrazábal M.E. 2009. Atividade diária de also be used in the defense of rest perches in Phalacrocorax Phalacrocorax brasilianus (Aves, Phalacrocoracidae), na região do carbo (Galván 2004). In the observed context, its function Saco da Fazenda, Itajaí, SC, Brasil. Ornithologia 3: 73‒82. would relate to the claim of the inanimate object acquired Casaux R., Ramón A., Tartara M.A., Borrell V. & Gonc R. 2012. Diet of breeding Neotropic Cormorants at the Carrileufu River, by another individual. Spot stealing, the posture assumed Patagonia: is there any impact on recreational fi sh resources? by one individual when climbing on the back of another Ardeola 59: 279‒289. and pecking its neck, refers to mating behavior (Berry Childress R.B. & Bennun L.A. 2002. Sexual character intensity and 1976). In addition, playing behaviors, such as looking its relationship to breeding timing, fecundity and mate choice in the Great Cormorant Phalacrocorax carbo lucidus. Journal of Avian for and manipulating leaves and branches, are similar to Biology 33: 23‒30. collecting material for the nest. Such behaviors among Chiquieri A., Soler M.F., Cassano M.L., Braz M.M. & Castro O. young individuals may favor the learning and practice of 1995. Vestibular. Rumos 10: 7‒28. innate responses, facilitating communication between Clark-Jr. G.A. 1969. Spread-wing postures in Pelecaniformes, adults (Sazima 2008). Ciconiiformes, and Falconiformes. Auk 86: 136‒139. Cook T.R. & Leblanc G. 2007. Why is wing-spreading behaviour In conclusion, this study provides important absent in Blue-eyed Shags? Animal Behaviour 74: 649‒652. information about the natural history of Neotropic D'Angelo G.B. & Sazima I. 2013. Commensal association of Cormorant and can be used as a starting point for further piscivorous birds with foraging otters in southeastern Brazil, and ethological studies on the species. a comparison with such a relationship of piscivorous birds with cormorants. Journal of Natural History 48: 241‒249. Espinoza-Rodríguez N., Carrasquero J., Turris-Morales K., Delgado- Ortega G. & Barrios-Garrido H. 2015. Asociaciones entre aves ACKNOWLEDGEMENTS marinas y Sotalia guianensis en el sur del Golfo de Venezuela. Caldasia 37: 309‒318. We thank anonymous reviewers for helping to improve Ferreira I., Ventura P.E.C. & Luz H.R. 2010. Aves no campus da Universidade Federal Rural do Rio de Janeiro. Rio de Janeiro: Edur. the manuscript and the National Council of Technological Galván I. 2004. Age-related spatial segregation of Great Cormorants and Scientifi c Development (Conselho Nacional de in a roost. Waterbirds 27: 377‒381. Desenvolvimento Científi co e Tecnológico – CNPq), for Gheler-Costa C., Comin F.H., Gilli L.C. & Verdade L.M. 2018. the scholarship awarded to T.L. da Silva during the Foraging behavior of Brazilian Cormorant, Nannopterum accomplishment of this work. brasilianus (Suliformes: Phalacrocoracidae). Zoologia 35: e14664. Gokula V. 2011. An ethogram of Spot-billed Pelican (Pelecanus philippensis). Chinese Birds 2: 183‒192. Gwiazda R. 2000. Numbers of non-breeding cormorants and their REFERENCES time budget activity at eutrophic, sub-mountain reservoir in southern Poland. 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Journal
Ornithology Research – Springer Journals
Published: Dec 1, 2018
Keywords: feeding; Lake Açu; maintenance; Phalacrocoracidae; seasonality