Estudos comportamentais de Paryphthimoides phronius (Lepidoptera: Satyrinae)

Orientador: Woodruff Whitman BensonDissertação (mestrado) - Universidade Estadual de Campinas, Instituto de BiologiaResumo: A evolução dos sistemas de acasalamento é uma área do comportamento animal intensamente estudada. Existe uma série de hipóteses que buscam estabelecer uma relação causal entre os diferentes sistemas de acasalamento e fatores ambientais que favorecem a sua origem e manutenção. Por exemplo, em espécies que naturalmente ocorrem em baixas densidades, os machos podem agregar em sítios específicos onde as chances de encontrar fêmeas são maiores. Em algumas situações, os machos podem defender territórios contra rivais para aumentar ainda mais a probabilidade de acasalamento. Algumas espécies de borboletas são fortemente territoriais e constituem bons modelos para o entendimento de como os fatores ambientais podem moldar os sistemas de acasalamento. Várias espécies temperadas da subfamília Satyrinae foram estudas, entretanto, trabalhos utilizando espécies tropicais são raros. Neste estudo, descrevi e testei algumas hipóteses acerca da origem, manutenção e organização dos sistemas de acasalamento em duas espécies desta subfamília: Paryphthimoides phronius e Hermeuptychia hermes. Em especial, demonstrei pela primeira vez que massa corporal é um importante fator relacionado com o sucesso territorial em uma borboleta e que esta característica prediz o sucesso territorial com maior precisão que o comprimento da asa. Também encontrei evidências de que frutas em decomposição podem ser importantes para o estabelecimento dos machos nos territórios. O trabalho foi organizado em três partes principais: 1) Identificação dos fatores climáticos que influenciam o padrão comportamental e a abundância durante o dia nas estações seca e chuvosa; 2) Descrição da organização do sistema de acasalamento e de sua variação anual com ênfase nas táticas dos machos de sua distribuição nos territórios; 3) Análise da natureza das interações entre machos, e de características fenotípicas que influenciem na probabilidade de vitória durante os confrontos. A defesa de território foi observada apenas P. phronius. A taxa de reavistamento de machos marcados foi de 34,9%, havendo indivíduos que apresentaram alta fidelidade nas áreas defendidas (variação menor que 9 metros entre os reavistamentos). Machos de P. phronius normalmente se estabelecem em manchas de sol no final da manhã, permanecendo nestas áreas até o entardecer. Esta espécie apresentou pico de abundância próximo ao meio dia, aumentando o número de indivíduos com a elevação da temperatura. Este resultado provavelmente está relacionado com a concentração de machos na borda da mata para defesa de territorial. O início da defesa coincidiu com o horário de nascimento das fêmeas, no entanto, fêmeas recém nascidas aparentemente não estão sexualmente receptivas. As áreas defendidas não estavam associadas com plantas hospedeiras, nem árvores em ambos os lados da estrada que possam favorecer a formação de manchas de sol. Hermeuptychia hermes apresentou baixa taxa de recaptura e ausência de defesa territorial. O padrão diário desta espécie foi caracterizado por um pico de abundância no início do dia e outro menor ao entardecer. Houve uma relação negativa entre abundância e temperatura apenas na estação chuvosa. Talvez H. hermes diminua a atividade nos horários mais quentes ou se desloque para o interior da mata. A ausência desta relação na estação seca pode ter sido causada por uma perda mais acentuada de calor que tenha impedido alterações comportamentais quando as temperaturas estavam maiores.Machos de P. phronius apresentam distribuição agregada ao longo dos trechos de estudo. Algumas parcelas foram recorrentemente ocupadas durante o ano, enquanto outras foram ocupadas somente nos períodos de alta densidade. Possivelmente os locais com freqüente ocupação de machos são mais valiosos para defesa. Por esta razão, estas áreas deveriam ser defendidas por machos com maior capacidade competitiva. Entretanto, não existiram diferenças nas características fenotípicas dos indivíduos que ocuparam estes segmentos e machos presentes em outros pontos. Observações preliminares sugerem que o estabelecimento de alguns machos está relacionado com a distribuição de frutas em decomposição. Para testar esta possibilidade frutas em decomposição foram colocadas em áreas em que estes insetos não ocorriam previamente. Este procedimento atraiu de 2 a 3 machos que se estabeleceram apenas nos sítios com manchas de sol. Os experimentos de remoção mostraram que os machos residentes foram, em média, 1,7 mg mais pesados que machos que os substituíram nos territórios, sugerindo que o aumento da massa corporal contribui para o sucesso territorial. Machos com maior comprimento de asa foram mais pesados com 25% da variação do peso sendo explicada pelo comprimento alar. Entretanto, a posse do território não estava associada com o comprimento das asas ou com seu desgaste (idade). Isto sugere que massa corporal está mais fortemente associada a fatores biológicos que determinam o RHP e que, medidas de comprimento de asa podem ser ineficientes como uma estimativa de tamanho, mesmo que o tamanho seja importante para a determinação do vencedor. Observações de disputas sugerem que machos mais pesados e com maior comprimento de asa não vencem mais disputas contra rivais mais leves e com menor comprimento alar. No entanto, o número de observações foi baixo. Machos com maior desgaste das asas (presumivelmente mais velhos) não foram mais leves que machos jovens. Indivíduos recapturados e pesados mais de uma vez mostraram uma tendência de redução do peso quando as recapturas tiveram um intervalo maior que três dias. As informações obtidas para P. phronius apresentaram algumas diferenças dos padrões observados em borboletas. A partir destes resultados sugiro algumas hipóteses e estudos posteriores para melhor esclarecer os mecanismos subjacentes aos comportamentos observados nesta espécieAbstract: The evolution of mating systems is an intensively studied area of animal behavior. There is a series of hypothesis establishing relationships between different mating systems and environmental factors that may favor their origin and maintenance. For example, in species that naturally occur in low densities males may aggregate at specific sites where the chances of encountering females are greater. In some situations, males may defend territories against rivals to increase even more their probability of mating. Some butterfly species are strongly territorial and constitute excellent systems for understanding how environmental factors mold mating systems. Studies using temperate species of the subfamily Satyrinae are abundant, however, the behavior of tropical species is poorly described. In this study, I have evaluated some hypothesis related to the origin, maintenance and organization of the mating systems of two Satyrinae; Paryphthimoides phronius and Hermeuptychia hermes. Especially, I show for the first time that body mass is an important correlate of territorial success in a butterfly and that it predict residence status much better than wing length. I also found evidence that decomposing fruit may be important in the shifts patterns of territory establishment. The study was divided in three main parts: 1) Identification of climatic factors that may influence adult behavior patterns and abundance during the day on winter and summer; 2) Description of mating system organization and its annual variation with particular reference to mating tactics and male distribution on territories; 3) Analysis of male agonistic interactions, and phenotypic characteristics related to contest success. Only P. phronius males exhibited territory defense. The male resighting rate was 34,9%, with some individuals showing high site fidelity in defended sites (variation smaller than 9 meters between resightings). Males normally defended sunspots between late morning and late afternoon. The abundance of Paryphtimoides phronius increased with temperature during the day, and was highest near midday. This increase during the hot periods is probably related to the concentration of males at edges for territorial defense. The time that territorial defense began coincided with the peak female emergence rate. Unexpectedly, newborn females did not seen to be sexually receptive. The defended sites were not associated with host plants or trees that may favor sunspots formation. Hermeuptychia hermes presented a low resighting rate and did not defend territories. The daily activity pattern of this species was characterized by an abundance peak in early morning and a smaller one in early afternoon. There was a negative relationship between abundance and temperature only in the raining season. Maybe H. hermes reduce activity in the hottest hours of the day or fly into the forest. The absence of this relationship during the dry season may have occurred due to a greater heat loss that prevented behavioral alterations when temperatures were higher. Paryphthimoides phronius showed an aggregated distribution and always defended territories near sunspots. Some plots were recurrently occupied during the year, while others were used only in periods of high density. The more frequently occupied sites may have been of higher quality, so Iexpected males with high competitive capacity at these points. However, no phenotypic differences existed between males defending more or less preferred sites. Preliminary observations suggest that males prefer territory sites near decomposing fruit. To test this possibility, fermented fruits were placed in areas previously unoccupied by P. phronius. This procedure attracted 2 or 3 males that set up territories at sunspots. Removal experiments showed that resident males were approximately 1,7 mg heavier than individuals that occupied the territory subsequently, suggesting that increased body mass contributed to territorial success. Heavier males also had longer wings with 25% of the variation in wing length explained by weight. However, ownership was not significantly related to wing length or wing wear (age) in this butterfly. This suggests that body mass is more closely related to the biological determinants of RHP and, even when size is important, wing length measures may be inefficient as a size estimative. Weight and forewing length did not predict the result of observed contests between males, although the sample size was small. More damaged males (presumably older) were not lighter in weight than younger ones. Nevertheless, mistakes during wing damage estimation may have occurred. Individuals recaptured and weighted more than once tended to lose weight for recaptures intervals longer than three days. The behaviors observed in P. phronius showed some differences in relation to the patterns observed in other butterflies. I suggest some hypothesis and posterior studies that may clarify the underlying mechanisms of the behaviors observed in this speciesMestradoEcologiaMestre em Ecologi

Cavity Length Affects the Occupation of Trap-Nests by Centris analis and Tetrapedia diversipes (Hymenoptera: Apidae)

The ideal cavity dimensions for neotropical cavity-nesting bees with the potential to be managed as pollinators have not been getting proper attention. We investigated whether the occupancy of trap-nests by Centris analis Fabricius and Tetrapedia diversipes Klug, and other nesting aspects, are affected by different trap-nest length. The used trap-nests were cardboard tubes 5, 10, 15 and 20 cm in length, and Ø 8 mm. Occupation rates of 10-cm trap-nests by C. analis was higher than that of the 5-cm ones (χ2=11.17, gl=1, p<0.001). On the other hand, there was not difference between the occupation rates of 10 and 15-cm long trap-nests (χ2=0.51, gl=1, p=0.48), and between the ones measuring 15 and 20 cm long (χ2=1.36, gl=1, p=0.24). T. diversipes occupied a smaller number of 5-cm trap-nests than the 10-cm ones (χ2=1.52, gl=1, p=0.22), as well as that the 15-cm ones were more occupied than the 10-cm trap-nests (χ2=4.23, gl=1, p=0.04); moreover, there was not difference between the occupation of 15 and 20-cm trap-nests (χ2=0.28, gl=1, p=0.59). Both species showed higher rates of dead immatures in nests set in the shortest trap-nests, whereas these mortality rates were lower in the longest ones. By taking into consideration that there was not significant difference in many of the assessed parameters in comparison to values recorded for 15 and 20-cm long trap-nests, it seems likely to recommend the adoption of 10-cm long trap-nests for C. analis reproduction in agricultural sites that depend on the pollination service provided by this bees species

Arboreal Ant Assemblages Respond Differently to Food Source and Vegetation Physiognomies: a Study in the Brazilian Atlantic Rain Forest

This study aimed to analyze assemblages of arboreal ants in different vegetation physiognomies within the Tropical Moist Forest (Atlantic Rain Forest) domain. The study was carried out at the Michelin Ecological Reserve, State of Bahia, Northeast of Brazil. We used sardine (protein resource) and honey (carbohydrate resource) baits to collect ants foraging in three vegetation types: (1) preserved native forest, (2) forest in regeneration (capoeira) with many invasive plants and (3) a mixed agroystem of rubber and cocoa tree plantation. We recorded 69 ant species attracted to the baits, 21 of them exclusive to honey bait and 25 exclusive to the sardine baits. The vegetation physiognomies preserved forest and rubber/cacao agrosystem showed higher species richness in relation to the forest in regeneration (capoeira), suggesting that rubber tree plantations can be a good matrix for the maintenance of some ant species typical of the forest matrix. The type of resource used is important for the structuring of the arboreal ant assemblages. The ants that were attracted to protein resources showed a guild composition that is more differentiated between vegetation types that of ants attracted to glucose resources

The Similar Usage of a Common Key Resource Does Not Determine Similar Responses by Species in A Community of Oil-collecting Bees

Variations in abundance and species richness among communities are often determined by interactions between biotic and abiotic factors. However, for communities composed of species that share a common specialization (such as similar foraging adaptations) it may be a key ecological factor involved in the common specialization that affects community variations. To evaluate this possibility, we characterized the guild of oil-collecting bees of a Neotropical savanna in Brazil and tested whether differences in Byrsonima abundance and availability of floral oil explain differences in species richness and abundance of oil-collecting bees of different tribes. Both the number of species and total abundance of Centridini species increased with the abundance of Byrsonima. One plausible explanation for the stronger adjustment between the abundance of Centridini and Byrsonima is that the abundance of these plants affects not only the availability of floral oil, but also of pollen. These findings indicate that the existence of a common specialization among different species does not homogenize their response to variations in a common explored resource

The role of parabiotic ants and environment on epiphyte composition and protection in ant gardens

Ant gardens (AGs) are a multi-partner specialized ant-plant interaction involving several ant and epiphyte species. Although studies on AGs have reported possible roles for some species in this system, there are unanswered questions regarding the process of epiphyte incorporation in the AGs and the role of less aggressive ant species in AG protection. In this study, we used AGs in the Brazilian Amazon forest formed by two parabiotic ant species to test a set of hypothesis regarding two main questions: 1) How is AG plant community composition affected by the surrounding environment? 2) Does Crematogaster levior play a role in the chemical detection of herbivory in the AGs? After identifying epiphytes occurring at AGs at the forest edge and in the interior, we found that ant gardens in each environment exhibited different compositions, and that plant species bearing oil or extrafloral nectar glands were more frequent in AGs located in the forest interior than in those at the forest edge. By performing experiments with volatile compounds emitted from injured epiphytes, we detected that only Camponotus femoratus was responsive, responding almost eight times faster in response to plant extracts than water treatments. Our results support the idea that environmental conditions affect ant preference for feeding resources provided by epiphytes and consequently shape the structure of the epiphyte community in AGs. On the other hand, the role of C. levior in AGs remains unknown, since it seems to play no direct or indirect role in AG protection

Daily Activity Patterns of Two Co-Occurring Tropical Satyrine Butterflies

Adult males and females of many insect species are expected to adjust their daily activity pattern in order to avoid stressful climatic conditions and increase the chances to encounter sexual partners. Using scan sampling methods associated with focal individual observations it was found that two satyrine butterflies of similar size and morphology, Hermeuptychia hermes (Fabricius) (Leptidoptera: Nymphalidae) and Paryphthimoides phronius (Godart), show completely different daily activity patterns on forest edges in southeastern Brazil. Hermeuptychia hermes presents one abundance peak in the morning and another in the late afternoon, while P. phronius abundance peaks in the mid-day, remaining stable until 1700 h. This difference is probably due to the occurrence of territorial behavior in the later species. The beginning of territorial defense by P. phronius males coincided with the time of new-born female activity. However, newly hatched females were not sexually receptive. The afternoon territoriality in male P. phronius may be in part related to mate acquisition. However, why the abundance of H. hermes decreases when the abundance of P. phronius increases is less clear

Pervasive gaps in Amazonian ecological research

Biodiversity loss is one of the main challenges of our time,1,2 and attempts to address it require a clear un derstanding of how ecological communities respond to environmental change across time and space.3,4 While the increasing availability of global databases on ecological communities has advanced our knowledge of biodiversity sensitivity to environmental changes,5–7 vast areas of the tropics remain understudied.8–11 In the American tropics, Amazonia stands out as the world’s most diverse rainforest and the primary source of Neotropical biodiversity,12 but it remains among the least known forests in America and is often underrepre sented in biodiversity databases.13–15 To worsen this situation, human-induced modifications16,17 may elim inate pieces of the Amazon’s biodiversity puzzle before we can use them to understand how ecological com munities are responding. To increase generalization and applicability of biodiversity knowledge,18,19 it is thus crucial to reduce biases in ecological research, particularly in regions projected to face the most pronounced environmental changes. We integrate ecological community metadata of 7,694 sampling sites for multiple or ganism groups in a machine learning model framework to map the research probability across the Brazilian Amazonia, while identifying the region’s vulnerability to environmental change. 15%–18% of the most ne glected areas in ecological research are expected to experience severe climate or land use changes by 2050. This means that unless we take immediate action, we will not be able to establish their current status, much less monitor how it is changing and what is being lostinfo:eu-repo/semantics/publishedVersio

Height and body-mass index trajectories of school-aged children and adolescents from 1985 to 2019 in 200 countries and territories: a pooled analysis of 2181 population-based studies with 65 million participants

Summary Background Comparable global data on health and nutrition of school-aged children and adolescents are scarce. We aimed to estimate age trajectories and time trends in mean height and mean body-mass index (BMI), which measures weight gain beyond what is expected from height gain, for school-aged children and adolescents. Methods For this pooled analysis, we used a database of cardiometabolic risk factors collated by the Non-Communicable Disease Risk Factor Collaboration. We applied a Bayesian hierarchical model to estimate trends from 1985 to 2019 in mean height and mean BMI in 1-year age groups for ages 5–19 years. The model allowed for non-linear changes over time in mean height and mean BMI and for non-linear changes with age of children and adolescents, including periods of rapid growth during adolescence. Findings We pooled data from 2181 population-based studies, with measurements of height and weight in 65 million participants in 200 countries and territories. In 2019, we estimated a difference of 20 cm or higher in mean height of 19-year-old adolescents between countries with the tallest populations (the Netherlands, Montenegro, Estonia, and Bosnia and Herzegovina for boys; and the Netherlands, Montenegro, Denmark, and Iceland for girls) and those with the shortest populations (Timor-Leste, Laos, Solomon Islands, and Papua New Guinea for boys; and Guatemala, Bangladesh, Nepal, and Timor-Leste for girls). In the same year, the difference between the highest mean BMI (in Pacific island countries, Kuwait, Bahrain, The Bahamas, Chile, the USA, and New Zealand for both boys and girls and in South Africa for girls) and lowest mean BMI (in India, Bangladesh, Timor-Leste, Ethiopia, and Chad for boys and girls; and in Japan and Romania for girls) was approximately 9–10 kg/m2. In some countries, children aged 5 years started with healthier height or BMI than the global median and, in some cases, as healthy as the best performing countries, but they became progressively less healthy compared with their comparators as they grew older by not growing as tall (eg, boys in Austria and Barbados, and girls in Belgium and Puerto Rico) or gaining too much weight for their height (eg, girls and boys in Kuwait, Bahrain, Fiji, Jamaica, and Mexico; and girls in South Africa and New Zealand). In other countries, growing children overtook the height of their comparators (eg, Latvia, Czech Republic, Morocco, and Iran) or curbed their weight gain (eg, Italy, France, and Croatia) in late childhood and adolescence. When changes in both height and BMI were considered, girls in South Korea, Vietnam, Saudi Arabia, Turkey, and some central Asian countries (eg, Armenia and Azerbaijan), and boys in central and western Europe (eg, Portugal, Denmark, Poland, and Montenegro) had the healthiest changes in anthropometric status over the past 3·5 decades because, compared with children and adolescents in other countries, they had a much larger gain in height than they did in BMI. The unhealthiest changes—gaining too little height, too much weight for their height compared with children in other countries, or both—occurred in many countries in sub-Saharan Africa, New Zealand, and the USA for boys and girls; in Malaysia and some Pacific island nations for boys; and in Mexico for girls. Interpretation The height and BMI trajectories over age and time of school-aged children and adolescents are highly variable across countries, which indicates heterogeneous nutritional quality and lifelong health advantages and risks

Worldwide trends in underweight and obesity from 1990 to 2022: a pooled analysis of 3663 population-representative studies with 222 million children, adolescents, and adults

Background Underweight and obesity are associated with adverse health outcomes throughout the life course. We estimated the individual and combined prevalence of underweight or thinness and obesity, and their changes, from 1990 to 2022 for adults and school-aged children and adolescents in 200 countries and territories. Methods We used data from 3663 population-based studies with 222 million participants that measured height and weight in representative samples of the general population. We used a Bayesian hierarchical model to estimate trends in the prevalence of different BMI categories, separately for adults (age ≥20 years) and school-aged children and adolescents (age 5–19 years), from 1990 to 2022 for 200 countries and territories. For adults, we report the individual and combined prevalence of underweight (BMI <18·5 kg/m2) and obesity (BMI ≥30 kg/m2). For schoolaged children and adolescents, we report thinness (BMI <2 SD below the median of the WHO growth reference) and obesity (BMI >2 SD above the median). Findings From 1990 to 2022, the combined prevalence of underweight and obesity in adults decreased in 11 countries (6%) for women and 17 (9%) for men with a posterior probability of at least 0·80 that the observed changes were true decreases. The combined prevalence increased in 162 countries (81%) for women and 140 countries (70%) for men with a posterior probability of at least 0·80. In 2022, the combined prevalence of underweight and obesity was highest in island nations in the Caribbean and Polynesia and Micronesia, and countries in the Middle East and north Africa. Obesity prevalence was higher than underweight with posterior probability of at least 0·80 in 177 countries (89%) for women and 145 (73%) for men in 2022, whereas the converse was true in 16 countries (8%) for women, and 39 (20%) for men. From 1990 to 2022, the combined prevalence of thinness and obesity decreased among girls in five countries (3%) and among boys in 15 countries (8%) with a posterior probability of at least 0·80, and increased among girls in 140 countries (70%) and boys in 137 countries (69%) with a posterior probability of at least 0·80. The countries with highest combined prevalence of thinness and obesity in school-aged children and adolescents in 2022 were in Polynesia and Micronesia and the Caribbean for both sexes, and Chile and Qatar for boys. Combined prevalence was also high in some countries in south Asia, such as India and Pakistan, where thinness remained prevalent despite having declined. In 2022, obesity in school-aged children and adolescents was more prevalent than thinness with a posterior probability of at least 0·80 among girls in 133 countries (67%) and boys in 125 countries (63%), whereas the converse was true in 35 countries (18%) and 42 countries (21%), respectively. In almost all countries for both adults and school-aged children and adolescents, the increases in double burden were driven by increases in obesity, and decreases in double burden by declining underweight or thinness. Interpretation The combined burden of underweight and obesity has increased in most countries, driven by an increase in obesity, while underweight and thinness remain prevalent in south Asia and parts of Africa. A healthy nutrition transition that enhances access to nutritious foods is needed to address the remaining burden of underweight while curbing and reversing the increase in obesit