Vocal and territorial behavior in the Smith frog (Hypsiboas faber): Relationships with plasma levels of corticosterone and testosterone

The possible trade-off between the roles of glucocorticoids as facilitators of energy substrate mobilization and neural inhibitors of sexual behavior during breeding season is under debate. We studied the relationship between calling and territorial behavior with plasma levels of corticosterone (CORT) and plasma levels of testosterone (T) across the breeding season of Hypsiboas faber, a large and territorial Neotropical treefrog. We investigated these relationships through focal observations of males calling naturally, followed by blood sampling for hormonal radioimmunoassay. We additionally used an experimental approach, which consisted of broadcasting recorded advertisement calls for 10 min to simulate an invasion in the territory of the focal subjects, followed by behavioral observation and blood sampling for hormonal radioimmunoassay. Results showed a pattern of co-variation between CORT and T across the breeding season. Furthermore, individual variation in CORT and T was related to different aspects of behavior: individuals with higher CORT showed higher calling rates, and individuals with higher steroid levels, mainly T, showed higher responsivity to social stimulation by other males in the chorus. Experimental simulation of territorial intrusion by using playback of advertisement calls of this species did not elicit consistent changes in agonistic behavior and CORT, but decreased T in focal males. (C) 2012 Elsevier Inc. All rights reserved.

Comportamento territorial de Hylodes nasus em um riacho na Floresta da Tijuca, Rio de Janeiro.

O comportamento territorial é exibido por animais que competem por recursos dentro de uma área específica defendida. Ao excluir competidores potenciais desta área através de comportamentos agressivos, os indivíduos territoriais garantem acesso prioritário aos recursos essenciais para sua sobrevivência e reprodução. Em anfíbios anuros, a territorialidade tem sido relacionada principalmente com espécies de reprodução prolongada, cujos sítios reprodutivos estão disponíveis durante todo o ano. O presente trabalho foi realizado em um trecho de riacho na Floresta da Tijuca, Rio de Janeiro. Foram realizadas quatro visitas consecutivas a área de amostragem em cada mês (abril/2009 à agosto/2010). O objetivo foi responder questões sobre a territorialidade dos indivíduos da espécie Hylodes nasus: (área de vida, fidelidade ao sítio, comportamentos agonísticos e combates físicos). As áreas de vida dos machos apresentaram relação significativa com o número de recapturas. O tamanho da área de vida das fêmeas foi ligeiramente maior do que dos machos. Os machos apresentaram maior fidelidade do que as fêmeas. A fidelidade apresentou resultado significativo com o CRC e a massa corporal dos indivíduos. Foram observadas 50 interações agonísticas entre machos, sendo exibidos 11 comportamentos agressivos. Foram observados 19 combates físicos, sendo os machos envolvidos classificados como residente, intruso, vencedor ou perdedor. Todas as disputas foram vencidas por indivíduos residentes. Informações a respeito da influência de fatores ambientais sobre a população de H. nasus, aspectos da estrutura populacional e descrição do microhábitat também são fornecidos.

Comportamento territorial de Lipaugus lanioides (Aves: Cotingidae) em área de Floresta Atlântica na Ilha Grande, RJ

O comportamento territorial é uma estratégia de monopolização de recursos quando esses são essenciais para o sucesso reprodutivo de um organismo. Um território é uma área de uso exclusivo, defendido contra invasores coespecíficos de mesmo sexo, resultante da interação social entre vizinhos. A territorialidade exerce importante papel no sistema reprodutivo de uma espécie, pois influencia a participação do macho na reprodução. Nesses casos, as fêmeas podem obter vantagens diretas, como sítios de nidificação e cuidado parental. O comportamento territorial também exerce influência na regulação do tamanho populacional através de uma relação entre custos e benefícios individuais: em ambientes ótimos e com alta densidade populacional, os territórios são pequenos com pouca substituição, e jovens machos têm dificuldade para conseguirem estabelecer-se. O presente estudo teve como objetivo investigar aspectos comportamentais do tropeiro-da-serra, espécie rara e endêmica de Floresta Atlântica, com distribuição bastante restrita. Ao longo de 18 meses na Ilha Grande (RJ), analisamos seu comportamento territorial, mensuramos os tamanhos de territórios individuais de machos e realizamos estimativas de densidade populacional. O playback foi utilizado para atestar a presença de territorialidade na espécie, para simular a aproximação de coespecíficos (interações intraespecíficas) e para induzir o deslocamento dos indivíduos até os limites de seus territórios. Para investigar as respostas comportamentais à aproximação de invasores, analisamos quantitativamente as reações dos indivíduos a estímulos sonoros (vocalização espontânea e induzida pelo playback). Os territórios individuais foram definidos em duas estações reprodutivas através do método do Mínimo Polígono Convexo (MPC) em uma área equivalente a 20ha. A densidade populacional foi definida através do número de territórios encontrados e pelo número de indivíduos vistos/ouvidos por unidade de área através de transecções lineares. As vocalizações espontâneas e induzidas ocorreram somente entre os meses de agosto a janeiro, caracterizando uma estação reprodutiva bem definida. Durante este período, os machos tornaram-se solitários e agressivos com coespecíficos; na fase não-reprodutiva, entretanto, os indivíduos mostram-se sociáveis, forrageando em pequenos grupos de até quatro indivíduos. Os resultados indicam que o território é estabelecido para a monopolização de alimento e acesso às fêmeas. Essas observações sugerem que a espécie estudada é territorialista. Foram estimados sete territórios com valores entre 0,21ha e 0,73ha (0,43 + 0,16ha). Os indivíduos apresentaram fidelidade territorial, ocupando os mesmos territórios em duas estações reprodutivas. A densidade populacional de L. lanioides apresentou flutuações ao longo do ano, com os maiores valores encontrados durante a estação reprodutiva (variando entre 0,37 e 1,84 indivíduos/ha). Flutuações na densidade populacional podem apontar migrações altitudinais motivadas por variações na disponibilidade de recursos alimentares. Concluímos que o comportamento reprodutivo de L. lanioides não se enquadra no conceito de sistema reprodutivo em leks, conhecido em outros cotingídeos (ex. Lipaugus vociferans), no qual a corte é um comportamento social, com disputa por status de dominância, e o papel do macho resume-se à cópula sem benefícios diretos para as fêmeas. Dessa forma, os resultados do presente estudo trazem informações originais sobre a biologia de L. lanioides

Behavior of the Amazonian damselfly Chalcopteryx scintillans McLachlan (Zygoptera: Polythoridae) and comments on its morphological distinction from C. rutilans (Rambur)

Coordenação de Aperfeiçoamento de Pessoal de Nível Superior (CAPES)

No scope for social modulation of steroid levels in a year-round territorial damselfish.

Both latitude and mating system have been proposed to shape relationships between steroid hormone levels and social behavior. Recently it has been postulated that species with long lasting non-seasonal territorial behavior have low androgen responsiveness. Tropical damselfishes are an ideal family to test this proposition because they show a large variety in mating systems. Here we contribute to the comparative dataset by measuring the response in steroid levels after social modulation in the banded sergeant, Abudefduf septemfasciatus, a species with non-seasonal territoriality. In highly territorial and brooding males, we found low androgen and cortisol levels that did not increase after experimental intraspecific simulated territorial intrusions (STI tests). No relationship was found between the variation in steroid hormone levels and territorial responses to naturally occurring territorial intrusions. Although steroid levels were low, male A. septemfasciatus were highly territorial both to STI challenges and to fishes that passed the territory. They often chased intruders for several meters away from the territory. This indicates that during nest defence in a non-seasonal territorial damselfish species, territorial behaviors are shown independent of variation in androgen and cortisol levels.

How resources control aggression in Drosophila

How animals use sensory information to weigh the risks vs. benefits of behavioral decisions remains poorly understood. Inter-male aggression is triggered when animals perceive both the presence of an appetitive resource, such as food or females, and of competing conspecific males. How such signals are detected and integrated to control the decision to fight is not clear. Here we use the vinegar fly, Drosophila melanogaster, to investigate the manner in which food and females promotes aggression.

In the first chapter, we explore how food controls aggression. As in many other species, food promotes aggression in flies, but it is not clear whether food increases aggression per se, or whether aggression is a secondary consequence of increased social interactions caused by aggregation of flies on food. Furthermore, nothing is known about how animals evaluate the quality and quantity of food in the context of competition. We show that food promotes aggression independently of any effect to increase the frequency of contact between males. Food increases aggression but not courtship between males, suggesting that the effect of food on aggression is specific. Next, we show that flies tune the level of aggression according to absolute amount of food rather than other parameters, such as area or concentration of food. Sucrose, a sugar molecule present in many fruits, is sufficient to promote aggression, and detection of sugar via gustatory receptor neurons is necessary for food-promoted aggression. Furthermore, we show that while food is necessary for aggression, too much food decreases aggression. Finally, we show that flies exhibit strategies consistent with a territorial strategy. These data suggest that flies use sweet-sensing gustatory information to guide their decision to fight over a limited quantity of a food resource.

Following up on the findings of the first chapter, we asked how the presence of a conspecific female resource promotes male-male aggression. In the absence of food, group-housed male flies, who normally do not fight even in the presence of food, fight in the presence of females. Unlike food, the presence of females strongly influences proximity between flies. Nevertheless, as group-housed flies do not fight even when they are in small chambers, it is unlikely that the presence of female indirectly increases aggression by first increasing proximity. Unlike food, the presence of females also leads to large increases in locomotion and in male-female courtship behaviors, suggesting that females may influence aggression as well as general arousal. Female cuticular hydrocarbons are required for this effect, as females that do not produce CH pheromones are unable to promote male-male aggression. In particular, 7,11-HD––a female-specific cuticular hydrocarbon pheromone critical for male-female courtship––is sufficient to mediate this effect when it is perfumed onto pheromone-deficient females or males. Recent studies showed that ppk23⁺ GRNs label two population of GRNs, one of which detects male cuticular hydrocarbons and another labeled by ppk23 and ppk25, which detects female cuticular hydrocarbons. I show that in particular, both of these GRNs control aggression, presumably via detection of female or male pheromones. To further investigate the ways in which these two classes of GRNs control aggression, I developed new genetic tools to independently test the male- and female-sensing GRNs. I show that ppk25-LexA and ppk25-GAL80 faithfully recapitulate the expression pattern of ppk25-GAL4 and label a subset of ppk23⁺ GRNs. These tools can be used in future studies to dissect the respective functions of male-sensing and female-sensing GRNs in male social behaviors.

Finally, in the last chapter, I discuss quantitative approaches to describe how varying quantities of food and females could control the level of aggression. Flies show an inverse-U shaped aggressive response to varying quantities of food and a flat aggressive response to varying quantities of females. I show how two simple game theoretic models, “prisoner’s dilemma” and “coordination game” could be used to describe the level of aggression we observe. These results suggest that flies may use strategic decision-making, using simple comparisons of costs and benefits.

In conclusion, male-male aggression in Drosophila is controlled by simple gustatory cues from food and females, which are detected by gustatory receptor neurons. Different quantities of resource cues lead to different levels of aggression, and flies show putative territorial behavior, suggesting that fly aggression is a highly strategic adaptive behavior. How these resource cues are integrated with male pheromone cues and give rise to this complex behavior is an interesting subject, which should keep researchers busy in the coming years.

Breeding of the Rufous-thighed Hawk (Accipiter erythronemius) in Argentina and Brazil

The Rufous-thighed, Hawk (Accipiter erythronemius) is a poorly-known, small raptor that dwells in forests and wooded savannas in south and eastern Brazil, Taraguay, eastern Bolivia, north and central Argentina, and Uruguay. We studied breeding biology and food habits of the Rufous-thighed Hawk from 1994 to 2007. We documented 14 breeding attempts recorded at nine nest sites in eight localities in Argentina and Brazil. Rufous-thighed Hawks were year-round residents at the study areas. The breeding season was c. 38 weeks long (June through March). Territorial behavior and courtship spanned for 22 weeks starting in mid June. Nest-building started in August. Nestlings were observed throughout December and into early January while fledglings were observed from late December through late March. Most nest sites were conifer plantations averaging 32.2 (SD = 10) years old, 12.6 ha (SD = 11.3) in size with trees averaging d 37.5 cm (SD = 13.5) of diameter at breast height. Nests were placed 20.2 m high (SD = 5.7), near the trunks of non-emergent conifers (Pinus sp. or Araucaria angustifolia). Nests were made of sticks, greater diameter averaging 48.8 cm (SD = 11.4), while nest depth averaged 23.3 cm (SD = 14). Prey items were birds (n = 49) ranging in size from c. 10 g [House Wren (Troglodytes, aedon)] to c. 140 g [White-tipped Dove (Leptotila verreauxi)]. Young started to chase birds at the age of 42-45 days in mid January. Adult males were primarily food providers but also aided in nest defense against other raptors. Females incubated, brooded, and fed young, and took leading roles in defense against humans. Rufous-thighed Hawks had unlined nests placed high in trees, and breeding season was markedly protracted.

Dietary L-tryptophan alters aggression in juvenile matrinxa Brycon amazonicus

This study evaluated the effect of dietary supplementation with -tryptophan (L-TRP), a serotonin precursor, on the aggressiveness of juvenile matrinx . Fish were kept in individual aquaria for 7 days receiving the diets: D1 (control: 0.47% of TRP), D2 (0.94% of TRP), D3 (1.88% of TRP), and D4 (3.76% of TRP). After this, they were grouped with an intruder fish to establish a resident-intruder relationship during periods of 20 min. Blood cortisol, glucose, chloride, sodium and calcium; hemoglobin, hematocrit, red blood cell count and volume; liver glycogen and lipids were measured. Territoriality had significant effect on the aggressiveness of matrinx (the residents were more aggressive than intruders, < 0.001) and tryptophan significantly affected their behavior. Fish fed with the D2 diet presented a longer latency until the first attack ( = 0.0069) and bit the intruder fewer times ( = 0.0136) during the period of observation, compared to the control group. The frequency of bites and chases after the first attack was not affected by the dietary supplementation of TRP. Physiological variables were not significantly affected by the diet, except for a moderate increase in cortisol level in fish fed with D2 diet after the fight, indicating slight activation of the hypothalamus-pituitary-interrenal axis. The results show that juvenile matrinx have aggressive and territorial behavior and that a diet containing 9.4 g TRP kg(-1) alter their aggressiveness, without affecting the stress-related physiological parameters.

Composição ictiofaunística e estratégias reprodutivas de quatro espécies de peixes nativos da Bacia Hidrográfica Piranhas - Assu, Rio Grande do Norte

This study focuses on the fish fauna composition and reproductive strategies of four native fish species in relation to environmental variables of the Piranhas-Assu hydrographic basin of the Caatinga biome, Rio Grande do Norte, Brazil. Fish and environmental data were collected monthly during the period September 2008 to February 2010, in the Marechal Dutra reservoir and River Acauã a tributary of Piranhas- Assu. The fish were measured, weighed and dissected, and the gonads were removed, weighed and examined macroscopically for identification and determination of gonad maturation stages. The results of this work generated seven articles. The first article is about the fish fauna composition of the Piranhas-Assu hydrographic basin, Rio Grande do Norte. The 602 fish samples captured were distributed in four orders (Characiformes, Perciformes, Siluriformes and Synbranchiformes), 11 families and 22 species, of which 17 are endemic to the Caatinga ecoregion. The order Characiformes was more representative followed by Perciformes, Siluriformes and Synbranchiformes. The second article is about the length-weight relationship and growth of seven native fish species Crenicichla menezesi, Cichlasoma orientale, Triportheus angulatus, Psectrogaster rhomboides, Pimelodella gracilis, Prochilodus brevis and Leporinus piau from a semiarid Brazilian reservoir. The third article is about the reproductive aspects of Crenicichla menezesi. Males were larger, heavier and with a slight predominance as compared to females. Four stages of gonadal development were characterized, being immature, maturing, mature and spent. The females reached sexual maturity earlier than males, with an average fecundity of 398 oocytes per batch. The spawning was partial with a long reproductive period. The fourth article deals with the reproductive strategy of Leporinus piau a neotropical freshwater fish in semi-arid region of Brazil. The population of L. piau (n = 211) showed a slight predominance of males (55%), with larger and heavier females. The males matured earlier than the females. This species presented total spawning, with an average fecundity of 55,000 mature oocytes. Rainfall and concentration of dissolved oxygen acted as influential factors during the spawning season. L. piau shows a seasonal reproductive strategy. The fifth article is related to the morphometric-meristic characteristics and reproductive aspects of freshwater sardine, Triportheus angulatus from River Acauã of the Caatinga biome. There was a predominance of larger females, reaching first maturation before males. There was total spawning during the rainy period of the region. The sixth article reports on the reproductive strategy of Psectrogaster rhomboides. The sex ratio was 1M: 1F, with negative allometric growth. Males reached sexual maturity earlier than females Females and males showed four stages of gonadal development and spawned during the rainy season. The fecundity was low and this species presented total spawning. The seventh article reports on the dynamics of territorial behavior of Crenicichla menezesi. Ten agonistic behaviors displayed by the males were observed: frontal and lateral threat, chasing, circular chasing, perpendicular, lateral and mouth attacks, escape, parallel positioning and stationary. The formation of four social groups was observed among males: without interaction; interaction with submission and escape; with frontal and lateral agonistic interactions. In social interactions between males and females, it was observed that the larger males interacted more among themselves and with the larger females. The large male established its territory and the two small males along with the small female were excluded from the other groups. These studies clarified the fish fauna composition and reproductive strategies of four native species Crenicichla menezesi, Leporinus piau, Triportheus angulatus and Psectrogaster rhomboides of the Piranhas- Assu hydrographic basin of the Caatinga biome, Rio Grande do Norte, Brazil.

Comportamento social e territorialidade alimentar na garça-azul, Egretta caerulea (L) = Feeding territoriality in the little blue heron, Egretta caerulea (L)

The use of habitat is an important part of a species biology. One resource of great importance for the survivor and reproduction of an individual is the food resource. Thus, the social interactions an animal has during the feeding activities are of extremely importance within its behavioral aspects, which represents the part of an organism trough which it interacts with the environment, adapting to changes and variations. Herons are known to form feeding aggregations of even more than thousands of individuals, in which social components of foraging have been identified and studied for several species. More profound studies of these aspects are yet to poor for the Little Blue Heron, Egretta caerulea. Therefore, the aim of this study was to describe the social behavior (display postures, vocalizations and co-specific interactions) and the territoriality of the specie during the feeding period in an area of mud bank in the estuarine system of Cananéia, south coast of São Paulo state, Brazil. The defense of a fixed and exclusive area, closest to the mangrove, trough expulsion was observed; some thing that have not yet been registered with concrete data for the specie. Higher capture and success rates, and lower investment rates (steps/min and stabs/min) were registered for individuals foraging in areas corresponding to the defended territory. This could be one of possible reasons for the establishment of territories in the area. Four display postures were registered for the specie, two of then new in the literature, which are used in the interactions between individuals; one vocalization, that apparently is important in the social context of foraging for the specie and, possibly, has a function of advertising and proclaiming the dominance position of the territorial individual within the group. A territorial individual uses three behaviors, of the ones described: expulsion, vocalization and encounter (agonistic encounter between individuals, without physical aggression). Of these, the expulsion is apparently used in the actual defense, actively; while the other two behaviors are used in a more passive way, in the maintenance of the dominance position of the individual, helping it in the defense of its territory in a less direct manner. Therefore, with the results presented in here, new components of the social utilization of the feeding resource for the Little Blue Heron were identified, incorporating aspects of the territorial behavior for a future understanding of its possible adaptive significance. And it also reinforces the importance of the social interactions of herons foraging in great aggregations, in areas ecologically important

Partilha de néctar de Eucalyptus spp., territorialidade e hierarquia de dominância em beija-flores (Aves: Trochilidae) no sudeste do Brasil

Territorial behavior in hummingbirds minimizes competition through aggressive interactions, resulting in a dominance hierarchy among species and individuals. Interactions among seven hummingbird species visiting flowering eucalyptus in the Floresta Estadual near Rio Claro, São Paulo, in southeastern Brazil, were studied. Dominance was determined by weight and size with the largest species being the most dominant. Time spent in defense and the number of aggressive interactions were greater than recorded in literature, perhaps due to the relatively greater density of hummingbirds in the study area. Daily activity patterns differed among dominant and subordinate species, but were not correlated with either the quantity of available nectar or with nectar sugar concentration.

The mesosomal gland of Xylocopa frontalis (Hymenoptera, Apidae, Xylocopini)

Males of Xylocopa fimbriata and X. micans present mesosomal glands that produce secretions containing many volatile substances with an intense floral fragrance, which resembles the fragrance produced by flowers that are attractive to the females of both species. The mating territories occupied by males of X. frontalis have no helpful resources for the females of this species, which are probably attracted to these environments by sexual pheromone(s) produced in the male mesosomal glands (lek polygyny). In this work, the presence of these glands in males of X. frontalis was confirmed and their anatomy and ultrastructure were described. An attempt to correlate the morphological characteristics of the mesosomal glands with the male territorial behavior was also carried out.

Sinalização visual e biologia reprodutiva de Dendropsophus werneri (Anura: Hylidae) em área de Mata Atlântica no estado do Paraná, Brasil

We studied the reproductive biology and visual signaling of Dendropsophus werneri, whose distribution is limited to the Atlantic Rain Forest. The fieldwork was carried out in the Estação 2 do IAPAR, municipality of Morretes, state of Paraná, Brazil, from August 2006 to March 2007. Additional information on reproduction was gathered in the Reserva Natural Salto Morato, municipality of Guaraqueçaba, state of Paraná, Brazil, from September 2006 to March 2007. Males were smaller than females. Males called throughout all the study period in Morretes, but were active only during three months in Guaraqueçaba; males called from low vegetation along the edge of temporary ponds in open areas. The visual signaling was observed in two contexts: (1) aggressive behaviors between two males and (2) during the reproduction, by amplectant males. In the territorial behavior, males exhibited both aggressive and mixed calls, visual signaling as well as physical combats. We also recorded satellite behavior in four males. The mean egg number per clutch was 244 ± 32 eggs, varying between 188 and 310 eggs. We observed two reproductive modes: Mode 1 and Mode 24. In the present study, Dendropsophus werneri showed elaborated social interactions involving visual signaling, territorial behavior, mating bahavior with tactile stimuli, and two different reproductive modes, demonstrating its complex reproductive biology.

Efeito da privação de ninho sobre a agressividade e o sucesso de acasalamento em machos de tilápia do Nilo

Conselho Nacional de Desenvolvimento Científico e Tecnológico (CNPq)

Área de vida da choca-do-planalto Thamnophilus pelzelni Hellmayr, 1924 (Passeriformes: Thamnophilidae) na estação ecológica de Jataí, Luiz Antônio, SP

Pós-graduação em Ciências Biológicas (Zoologia) - IBRC