Establishment and Expansion Scenario of Drosophila suzukii (Diptera: Drosophilidae) in Central Brazil.

Drosophila suzukii Matsumura (Diptera: Drosophilidae), also known as spotted wing drosophila (SWD), is an important pest that damages various wild and cultivated soft fruits worldwide, especially in the Northern Hemisphere. In Brazil, it occurs mainly in the subtropical climates of the southern and southeastern regions. However, SWD has also been sporadically found in the central region of the country in the natural vegetation of the tropical Brazilian Savanna. In this study, we investigated the occurrence of SWD at the northern limit of its range in South America - the central region of Brazil - by monitoring an established drosophilid community in an orchard located in the Brazilian Federal District. We also investigated the current geographical distribution of this pest in Brazil and its potential geographical distribution using species distribution models under two different future shared socioeconomic pathways scenarios (2040 and 2060, optimist and pessimist). Twenty drosophilid species were detected among the 6,396 captured specimens, most of which are exotic in the Neotropical region. The fly community greatly fluctuated throughout the year, and the highest abundance of SWD (3.5% relative abundance and 1.38 flies/trap/day) was recorded in April during the rainy season. Potential distribution models indicate that suitable areas for SWD spread will decrease in the south and southeast but increase in the central region of Brazil. We recommend continuous SWD monitoring and improving bioclimatic forecast models for mitigating damage to local fruit production.

Host Preferences Shown by Drosophilids (Diptera) in a Commercial Fruit and Vegetable Distribution Center Follow the Wild Neotropical Pattern.

Although drosophilids have been extensively studied in laboratories worldwide, their ecology is still relatively poorly understood. This is unfortunate because some species are currently expanding their geographic distribution and infesting fruit crops. Here, we investigated the relationship between drosophilids and potential plant hosts in a commercial fruit and vegetable distribution center in the Neotropical region. We collected discarded fruits and vegetables from this commercial center during two time periods (2007-2008 and 2017-2018). Resources were weighted and individually monitored in the laboratory. The drosophilids that emerged were identified, and the relationship between them and their resources was explored. From the 99,478 kg of potential hosts collected, we identified 48 plant taxa, from which 48,894 drosophilids of 16 species emerged. On both collecting occasions, drosophilid assemblages were strongly dominated by basically the same exotic species, which explore a broader range of resources, especially those of exotic origin, when compared to neotropical drosophilids. These results are concerning because the studied site, Along with other urban markets around the world, might be acting as sources of generalist widespread species that disperse to surrounding natural vegetation and contribute to biotic homogenization.

Parasitoids of Drosophilids in the Brazilian Savanna: Spatial-temporal Distribution and Host Associations with Native and Exotic Species.

Host-parasitoid interactions may have a relevant role not only in ecological processes but also in human procedures such as biological control and the management of invasive species. Although the Drosophila-parasitoid system has been widely used for investigating all aspects of host-parasitoid relationships, it is still poorly understood in tropical areas. Here, we investigate the richness, spatial-temporal distribution, and host associations of parasitoids attacking drosophilid flies in the core region of the Brazilian Savanna, a hotspot of biodiversity. Using different methods, we collected wasps on several occasions over 7 years and found 13 species representing families associated with drosophilid larvae (Figitidae, Braconidae) and pupae (Pteromalidae, Eurytomidae, Dipriidae). The dominant species infesting larvae and pupae were Dicerataspis grenadensis Ashmead and Pachycrepoideus vindemmiae (Rondani), respectively. Spalangia simplex Perkins was recorded for the first time in the Brazilian Savanna. Although our study was not designed to evaluate collection methods, we observed that they captured different subsets of the wasp community. The spatiotemporal distribution of wasps followed those of drosophilids: they were more abundant in forests and during the rainy season, suggesting that the abundance of hosts, especially native drosophilids, is an important factor determining the assemblage structure and population dynamics of parasitoids. Common parasitoids and drosophilids were associated with more than one host/antagonist species, suggesting that caution should be used for the selection of biological control agents. This study confirms the role and relevance of natural vegetation in preserving biodiversity and ecosystem services, especially in a biome severely threatened by agricultural expansion.

Identification key for drosophilid species (Diptera, Drosophilidae) exotic to the Neotropical Region and occurring in Brazil

Abstract Thirteen species of drosophilid exotic to the Neotropical Region are recorded in Brazil, and some of them are highly invasive and threaten significantly fruit cultures. We provide an illustrated key for identifying these species, and briefly discuss their taxonomic status, distribution, and occurrence in the Neotropics. The key should not only support newcomers to the study of Drosophila but also facilitate their identification by those interested in insects associated with cultivated areas.

Seasonal variation of a plant-pollinator network in the Brazilian Cerrado: Implications for community structure and robustness.

Seasonal variation in the availability of floral hosts or pollinators is a key factor influencing diversity in plant-pollinator communities. In seasonally dry Neotropical habitats, where month-long periods of extreme drought are followed by a long rainy season, flowering is often synchronized with the beginning of precipitation, when environmental conditions are most beneficial for plant reproduction. In the Brazilian Cerrado, a seasonally dry ecosystem considered one of the world's biodiversity hotspots for angiosperms, plants with shallow root systems flower predominantly during the rainy season. Foraging activity in social bees however, the major pollinators in this biome, is not restricted to any particular season because a constant supply of resources is necessary to sustain their perennial colonies. Despite the Cerrado's importance as a center of plant diversity, the influence of its extreme cycles of drought and precipitation on the dynamics and stability of plant-pollinator communities is not well understood. We sampled plant-pollinator interactions of a Cerrado community weekly for one year and used network analyses to characterize intra-annual seasonal variation in community structure. We also compared seasonal differences in community robustness to species loss by simulating extinctions of plants and pollinators. We find that the community shrinks significantly in size during the dry season, becoming more vulnerable to disturbance due to the smaller pool of floral hosts available to pollinators during this period. Major changes in plant species composition but not in pollinators has led to high levels of turnover in plant-pollinator associations across seasons, indicated by in interaction dissimilarity (<3% of shared interactions). Aseasonal pollinators, which mainly include social bees and some solitary specialized bees, functioned as keystone species, maintaining robustness during periods of drastic changes in climatic conditions.

Sexual size dimorphism in three species of forensically important blowflies (Diptera: Calliphoridae) and its implications for postmortem interval estimation.

Forensic entomologists rely on insects present in human remains to establish a minimum postmortem interval (PMImin). Blowflies have been widely used in these estimates because they lay eggs on the victim's body shortly after death and, within hours, larvae hatch and grow at predictable rates. Hence, isomegalen diagrams based on larval size and local temperature are considered good models to estimate PMImin. Still, most professional do not account for size sexual dimorphism in blowflies, which add uncertainty to predictive models by two mechanisms: (1) males and females might grow up to different sizes, and; (2) males and females might grow at different rates. In this study, we investigate sexual dimorphism and biases on the prediction of adult size in three species of blowflies (Lucilia sericata, Calliphora vicina and Calliphora vomitoria) reared under different larval densities. Estimated size range, stablished with and without sex discrimination, showed that females were larger than males in the three species. The ultimate size of adult stage, however, was more difficult to predict. C. vicina and C. vomitoria decrease as density raises but at different rates, and even males and females of the same species react differently to density increase. Adult size of L. sericata, in contrast, shows a slight increase with density. Except for C. vomitoria females, estimated size ranges are lower when species are divided by sex. Our results show that sex is an important factor to consider in PMImin estimates. Scenarios for all three species shorten their estimated size ranges when compared to databases with no sex identification. Therefore, computing data by sex raises accuracy in size based predictive models.

Differential attraction of drosophilids to banana baits inoculated with Saccharomyces cerevisiae and Hanseniaspora uvarum within a Neotropical forest remnant.

BACKGROUND: Yeasts are a necessary requisite in the diet of most Drosophila species that, in turn, may vector their dispersal in natural environments. Differential attractiveness experiments and the isolation of yeasts consumed by Drosophila may be informative for characterizing this association. Hanseniaspora uvarum is among the most common yeast species isolated from Drosophila crops, with high attractiveness to drosophilids. Saccharomyces cerevisiae has been widely used to collect flies, and it allows broad sampling of almost all local Drosophila species. Pronounced differences in the field concerning Drosophila attractivity to baits seeded with these yeast species have been previously reported. However, few explicit generalizations have been set. Since late fifties, no field experiments of Drosophila attractivity were carried out in the Neotropical region, which is facing shifts in abiotic and biotic factors. Our objective is to characterize preference behavior that mediates the interaction in the wild among Neotropical Drosophila species and yeasts associated with them. We want to set a broad generalization about drosophilids attracted to these yeasts. Here we present the results of a differential attractiveness experiment we carried out in a natural Atlantic Rainforest fragment to assess the preferences of Drosophila species groups to baits inoculated with H. uvarum and S. cerevisiae. METHODS: Both yeast species were cultured in GYMP broth and separately poured in autoclaved mashed banana that was left fermenting. In the field, we collected drosophilids over five arrays of three different baits: non-inoculated autoclaved banana and banana inoculated with each yeast. In the laboratory the drosophilids were sorted to five sets according to their external morphology and/or genitalia: tripunctata; guarani; willistoni; exotic; and the remaining flies pooled in others. RESULTS AND CONCLUSIONS: Uninoculated banana baits attracted virtually no flies. We found significant departures from random distribution over the other two baits (1:1 proportion) for all sets, except the pooled others. Flies of the sets willistoni and exotic preferred H. uvarum over S. cerevisiae, while the remaining sets were more attracted to S. cerevisiae. Previously, various authors reported similar patterns in attraction experiments with S. cerevisiae and H. uvarum. It is also noteworthy that both yeast species have been isolated from natural substrates and crops of Drosophila species. Taken together, these results suggest that the preferences among Drosophila species groups may be reflecting deep and stable relations with yeast species in natural environments. They can be summarized as: forest dwelling species from subgenus Drosophila (such as tripunctata and guarani groups) are attracted to banana baits seeded with S. cerevisiae; while exotic (as D. melanogaster) and subgenus Sophophora species are preferentially attracted to baits seeded with H. uvarum.

Ecological insights from assessments of phenotypic plasticity in a Neotropical species of Drosophila.

Several authors have called attention to the evolutionary importance of phenotypic plasticity and niche construction, because such phenomena require a new status and a new perspective. Drosophila species are traditionally used as models in investigations of phenotypic plasticity, although the majority of such research has been conducted with species of the subgenus Sophophora, primarily Drosophila melanogaster. In this study, we investigated the phenotypic plasticity of Drosophila cardini, a Neotropical species of the subgenus Drosophila, and focused on the wing size, wing shape, thorax length and wing: thorax ratio of lines that were collected in the Brazilian savanna and exposed to different temperatures during growth. All of the analyzed traits presented plasticity to temperature, and the reaction norms were similar to those previously found in other drosophilid species; in addition, the maximum values were consistent with the temperature variations at the collection sites. The specimens that emerged at low temperatures were larger and had more rounded wings compared with those that emerged at high temperatures, which were smaller and had narrower wings. We hypothesized that the differences observed in the shape of the wings might be associated with flight performance. Nevertheless, further investigation of the relationships among wing shape, wing loading and flight performance is required. Investigations on phenotypic plasticity using species with diverse ecologies should help us to better understand how this phenomenon operates in nature, and studies of this type must be encouraged.

FAMILY DROSOPHILIDAE.

This catalogue presents 176 species distributed in 17 genera and four tribes of Drosophilidae in Colombia, with both subfamilies, Drosophilinae and Steganinae, represented in the Colombian territory. This is probably an underestimate of the real richness of drosophilid species in the country, for two reasons. First, there are relatively few collections of Drosophilidae in Colombia, resulting in significant spatial gaps in the sampling of these flies. Second, we have not considered here species that have already been recorded in neighbour countries but not in Colombia. Consequently, there are certainly several described species, as well as new species, that occur in this area but are not listed here. We hope that this catalogue will stimulate new inventories of drosophilids in Colombia, contributing to fill gaps in the knowledge of this family of flies in the country.

Spatial and temporal dynamics of drosophilid larval assemblages associated to fruits

The study of organisms and their resources is critical to further understanding population dynamics in space and time. Although drosophilids have been widely used as biological models, their relationship with breeding and feeding sites has received little attention. Here, we investigate drosophilids breeding in fruits in the Brazilian Savanna, in two contrasting vegetation types, throughout 16 months. Specifically, larval assemblages were compared between savannas and forests, as well as between rainy and dry seasons. The relationships between resource availability and drosophilid abundance and richness were also tested. The community (4,022 drosophilids of 23 species and 2,496 fruits of 57 plant taxa) varied widely in space and time. Drosophilid assemblages experienced a strong bottleneck during the dry season, decreasing to only 0.5% of the abundance of the rainy season. Additionally, savannas displayed lower richness and higher abundance than the forests, and were dominated by exotic species. Both differences in larval assemblages throughout the year and between savannas and gallery forests are consistent with those previously seen in adults. Although the causes of this dynamic are clearly multifactorial, resource availability (richness and abundance of rotten fruits) was a good predictor of the fly assemblage structure.

Pterandra pyroidea: a case of pollination shift within neotropical Malpighiaceae.

BACKGROUND AND AIMS: Most Neotropical species of Malpighiaceae produce floral fatty oils in calyx glands to attract pollinating oil-collecting bees, which depend on this resource for reproduction. This specialized type of pollination system tends to be lost in members of the family that occur outside the geographic distribution (e.g. Africa) of Neotropical oil-collecting bees. This study focused on the pollination ecology, chemical ecology and reproductive biology of an oil flower species, Pterandra pyroidea (Malpighiaceae) from the Brazilian Cerrado. Populations of this species consist of plants with oil-secreting (glandular) flowers, plants with non-oil-secreting flowers (eglandular) or a mix of both plant types. This study specifically aims to clarify the role of eglandular morphs in this species. METHODS: Data on pollinators were recorded by in situ observations. Breeding system experiments were conducted by isolating inflorescences and by enzymatic reactions. Floral resources, pollen and floral oils offered by this species were analysed by staining and a combination of various spectroscopic methods. KEY RESULTS: Eglandular flowers of P. pyroidea do not act as mimics of their oil-producing conspecifics to attract pollinators. Instead, both oil-producing and oil-free flowers depend on pollen-collecting bees for reproduction, and their main pollinators are bumble-bees. Floral oils produced by glandular flowers are less complex than those described in closely related genera. CONCLUSIONS: Eglandular flowers represent a shift in the pollination system in which oil is being lost and pollen is becoming the main reward of P. pyroidea flowers. Pollination shifts of this kind have hitherto not been demonstrated empirically within Neotropical Malpighiaceae and this species exhibits an unusual transition from a specialized towards a generalized pollination system in an area considered the hotspot of oil-collecting bee diversity in the Neotropics. Transitions of this type provide an opportunity to study ongoing evolutionary mechanisms that promote the persistence of species previously involved in specialized mutualistic relationships.

Breeding sites of drosophilids (Diptera) in the Brazilian Savanna. I. Fallen fruits of Emmotum nitens (Icacinaceae), Hancornia speciosa (Apocynaceae) and Anacardium humile (Anacardiaceae) / Sítios de criação de drosofilídeos (Diptera) no bioma Cerrado. I. Frutos caídos de Emmotum nitens (Icacinaceae), Hancornia speciosa (Apocynaceae) e Anacardium humile (Anacardiaceae)

In this study, drosophilids that bred in fruits of three common plant species of the Brazilian Savanna were investigated: Emmotum nitens, Hancornia speciosa and Anacardium humile, along with the temporal and spatial distribution of these insects among fruits obtained from six individuals of E. nitens. Fallen fruits were collected in natural environments, placed on moist sand in individual containers and all drosophilids that emerged from these resources during 15 days were collected, counted and identified. From 3,651 fruits collected (3,435 of E. nitens, 179 of A. humile and 37 of H. speciosa) 4,541 flies emerged and were classified into 19 species of Drosophilidae. Their distribution was unequal among the three resources, also over time, and among the six individuals of E. nitens. Such fluctuations probably reflect the availability of resources in time and space and probably the action of selective factors such as larval competition.

Neste estudo, são investigados os drosofilídeos que se criam em frutos de três espécies de plantas comuns do Cerrado brasileiro: Emmotum nitens, Hancornia speciosa e Anacardium humile, juntamente com a distribuição temporal e espacial desses insetos entre os frutos obtidos de seis indivíduos de E. nitens. Frutos recolhidos sobre a serrapilheira, em ambientes naturais, foram mantidos no laboratório e os drosofilídeos que deles emergiram foram contados e identificados. Dos 3.651 frutos coletados (3.435 de E. nitens, 179 de A. humile e 37 de H. speciosa) emergiram 4.541 drosofilídeos, classificados em 19 espécies. A distribuição dessas espécies foi desigual entre os três tipos de recursos, ao longo do tempo, e também entre os seis indivíduos de E. nitens. Essas flutuações refletem a disponibilidade dos recursos no tempo e no espaço, e provavelmente a ação de forças seletivas como a competição entre as larvas.

Biogeographical aspects of drosophilids (Diptera, Drosophilidae) of the Brazilian savanna / Aspectos biogeográficos dos drosofilídeos (Diptera, Drosophilidae) da savana brasileira

Some species of drosophilids have been intensely studied, especially in the areas of genetics and molecular biology. However, little is known about their ecology and biogeography. Here, we revise the records of Drosophilidae species in the Cerrado domain, in order to organize the available information and to recognize patterns of geographic distribution and endemism. The 43 sampled sites in this domain reveal 100 species of drosophilids (10 of them exotic to the Neotropical region), distributed in eight genera. The collections were concentrated mainly in the States of Goiás (including the Federal District) and Minas Gerais. Serra do Cipó is the site with the highest richness of these insects. Of the 90 neotropical species found in the Cerrado, 77 also occur outside this domain, most of which are in the Atlantic Forest. Since extensive areas of this domain were poorly sampled for drosophilids and most studies used only baited traps to collect the flies, this absence of endemism should be carefully considered. It is recommended new inventories in poorly sampled regions, especially in the northeastern part of the domain. Also diversification of collection methods and improvement in the taxonomy of these flies are proposed. These directions should provide a significant increase in the list of species of drosophilids of the Cerrado; and thus, increase our knowledge about its biodiversity.

Algumas espécies de drosofilídeos têm sido intensamente estudadas, principalmente nas áreas de genética e biologia molecular. Entretanto, a ecologia e biogeografia do grupo ainda são pouco conhecidas. Aqui revisamos os registros das espécies de Drosophilidae no domínio do Cerrado, visando organizar a informação disponível e reconhecer eventuais padrões de distribuição geográfica e endemismos. As 43 localidades amostradas no domínio apresentaram 100 espécies de drosofilídeos (10 delas exóticas à região Neotropical), distribuídas em oito gêneros. As coletas estão concentradas principalmente nos Estados de Goiás (incluindo Distrito Federal) e Minas Gerais. A Serra do Cipó é o local com maior riqueza desses insetos. Das 90 espécies neotropicais encontradas no Cerrado, 77 também ocorrem fora do domínio, a maioria na Mata Atlântica. Como áreas extensas do domínio foram pouco amostradas para drosofilídeos e a maior parte dos estudos utilizaram apenas iscas de banana para coletar as moscas, a ausência de endemismos deve ser analisada com precaução. São recomendados novos inventários em regiões pobremente amostradas, especialmente na parte noroeste do domínio. Também são propostas a diversificação dos métodos de coleta e o aumento do esforço de identificação taxonômico dessas moscas. Essas diretrizes devem permitir um aumento significativo na lista de espécies de drosofilídeos do Cerrado e, consequentemente, no conhecimento da biodiversidade do domínio.

Drosophilids (Insecta, Diptera) of the Paranã Valley: eight new records for the Cerrado biome / Drosofilídeos do Vale do Paranã: oito novas ocorrências para o bioma Cerrado

The Paranã Valley, located in the Central Brazil Plateau, within the domain of the Cerrado biome, is frequently cited as a center of endemism and diversity. Nonetheless, this region is poorly known, mainly considering its invertebrate fauna. Here, drosophilid flies were used as biological tools to contribute to a broader inventory, whose goals were mapping the Cerrado biodiversity and identifying areas to be conserved. Three previously unsampled areas in the Paranã Valley were sampled. Amongst the 12,297 specimens collected 45 species were identified, which represents 50 percent of the drosophilid species previously recorded in the Cerrado. Moreover, this sampling presented eight new occurrences for the biome (Drosophila annulosa, D. calloptera, D. papei, D. neomorpha, D. roehrae, Gitona bivisualis, Rhinoleucophenga lopesi, and the genus Diathoneura) and three species that had only one record for the biome, or that were recorded only in a specific area. Therefore, this study confirms the importance of the Paranã Valley as a center of biodiversity in the Cerrado biome, and recommends the establishment of conservation units in this region.

O Vale do Paranã localiza-se no Brasil Central, no domínio do bioma Cerrado, e é freqüentemente citado como um centro de diversidade e endemismo. Entretanto, essa região ainda é pouco conhecida, principalmente no que se refere à fauna de invertebrados. Este estudo apresenta a fauna de drosofilídeos de três áreas nunca avaliadas anteriormente no Vale do Paranã, visando contribuir para um inventário mais amplo, cujos objetivos foram mapear a biodiversidade da região e identificar áreas para serem preservadas. Dentre os 12.297 espécimes coletados foram identificadas 45 espécies, correspondendo a 50 por cento dos drosofilídeos previamente registrados no Cerrado. Além disso, este trabalho acrescentou oito novas ocorrências para o Cerrado (Drosophila annulosa, D. calloptera, D. papei, D. neomorpha, D. roehrae, Gitona bivisualis, Rhinoleucophenga lopesi e o gênero Diathoneura) e três outras espécies com apenas uma única ocorrência para o Cerrado, ou registradas em apenas uma área específica do bioma. Nesse contexto, este estudo confirma a importância do Vale do Paranã como um centro de biodiversidade do Cerrado, e recomenda o estabelecimento de unidades de conservação nessa região.

Learning of courtship components in Drosophila mercatorum (Paterson & Wheller) (Diptera, Drosophilidae)

In Drosophila, courtship is an elaborate sequence of behavioural patterns that enables the flies to identify conspecific mates from those of closely related species. This is important because drosophilids usually gather in feeding sites, where males of various species court females vigorously. We investigated the effects of previous experience on D. mercatorum courtship, by testing if virgin males learn to improve their courtship by observing other flies (social learning), or by adjusting their pre-existent behaviour based on previous experiences (facilitation). Behaviours recorded in a controlled environment were courtship latency, courtship (orientation, tapping and wing vibration), mating and other behaviours not related to sexual activities. This study demonstrated that males of D. mercatorum were capable of improving their mating ability based on prior experiences, but they had no social learning on the development of courtship.

Em Drosophila, a corte sexual consiste em uma elaborada seqüência de padrões comportamentais que possibilita às moscas reconhecer parceiros conspecíficos dentre indivíduos de outras espécies. Essa discriminação é importante uma vez que drosofilídeos geralmente se agregam em sítios de alimentação, onde machos de diversas espécies cortejam as fêmeas vigorosamente. Neste estudo, testamos se machos virgens aprimoram seu comportamento de corte mediante a observação do cortejo de outras moscas da população (aprendizado social), ou mediante experiências próprias anteriores (facilitação). O comportamento de corte das moscas, observado em ambiente controlado, foi registrado com relação aos seguintes componentes: latência da corte, duração da corte (orientação, "tapping" e vibração das asas), cópula, e outros comportamentos não relacionados à corte sexual. Os resultados aqui obtidos sugerem que machos de D. mercatorum são capazes de aprimorar sua habilidade de cortejar fêmeas em função de experiências próprias anteriores, mas não encontramos nenhuma evidência de aprendizado social para esse comportamento.

Teaching evolutionary biology

Evolutionary Biology integrates several disciplines of Biology in a complex and interactive manner, where a deep understanding of the subject demands knowledge in diverse areas. Since this knowledge is often inaccessible to the majority of specialized professionals, including the teachers, we present some reflections in order to stimulate discussions aimed at the improvement of the conditions of education in this area. We examine the profile of evolutionary teaching in Brazil, based on questionnaires distributed to teachers in Secondary Education in the Federal District, on data provided by the "National Institute for Educational Studies and Research", and on information collected from teachers working in various regions of this country. Issues related to biological misconceptions, curriculum and didactic material are discussed, and some proposals are presented with the objective of aiding discussions aimed at the improvement of the teaching of evolutionary biology.