515 resultados para FICUS CARICA


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Mutualisms are interspecific interactions in which both players benefit. Explaining their maintenance is problematic, because cheaters should outcompete cooperative conspecifics, leading to mutualism instability. Monoecious figs (Ficus) are pollinated by host-specific wasps (Agaonidae), whose larvae gall ovules in their "fruits'' (syconia). Female pollinating wasps oviposit directly into Ficus ovules from inside the receptive syconium. Across Ficus species, there is a widely documented segregation of pollinator galls in inner ovules and seeds in outer ovules. This pattern suggests that wasps avoid, or are prevented from ovipositing into, outer ovules, and this results in mutualism stability. However, the mechanisms preventing wasps from exploiting outer ovules remain unknown. We report that in Ficus rubiginosa, offspring in outer ovules are vulnerable to attack by parasitic wasps that oviposit from outside the syconium. Parasitism risk decreases towards the centre of the syconium, where inner ovules provide enemy-free space for pollinator offspring. We suggest that the resulting gradient in offspring viability is likely to contribute to selection on pollinators to avoid outer ovules, and by forcing wasps to focus on a subset of ovules, reduces their galling rates. This previously unidentified mechanism may therefore contribute to mutualism persistence independent of additional factors that invoke plant defences against pollinator oviposition, or physiological constraints on pollinators that prevent oviposition in all available ovules.

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In this preliminary study, the reproductive phenology of two monoecious fig species, Ficus racemosa and F. rubiginosa, was examined in tropical Australia. Syconia (inflorescences) occurred on both species all year round, but pre-floral and interfloral syconia were much commoner than the wasp-receptive and wasp-emitting phases in both species. The temporal overlap of the wasp-receptive and wasp-emitting phases on a single tree indicated that self-pollination was possible in both species and that pollinators may sometimes persist through multiple generations on one tree. This sexual phase overlap was commoner in F. rubiginosa than in F racemosa. The two species also differed in their general within-tree asynchrony, with a higher diversity of phases on F. rubiginosa than on F. racemosa. The time from syconium initiation to ripening was very similar in F. rubiginosa (mean = 48.51 days) and F. racemosa (mean = 43.53 days). However, there was much more variation within and between trees for F. rubiginosa. In addition, the wasp-receptive phase was found to last up to 5 days (rnean = 4.38) in F. rubiginosa. Such longevity should contribute substantially to local pollinator population persistence. Future work should use genetic studies to determine whether self-pollination is common in these fig species.

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Although theory exists concerning the types of strategies that should be used in contests over resources, empirical work explicitly testing its predictions is relatively rare. We investigated male fighting strategies in two nonpollinating. g wasp species associated with Ficus rubiginosa figs. In Sycoscapter sp. A, males did not assess each other before or during fights over mating opportunities. Instead,fights continued until the loser reached an energetic cost threshold that was positively correlated with its body size (fighting ability) and retreated. In Philotrypesis sp. B, pre fight assessment was indicated, with males attacking competitively inferior rivals to remove them from the competitor pool ( they then continued to do so until they reached a cost threshold that was again positively correlated with body size). Using data on species ecology, we discuss our findings with respect to theory on when different fighting strategies should evolve. We argue that the type of strategy used by a. g wasp species is determined by its relative benefits in terms of inclusive fitness. (c) 2008 The Association for the Study of Animal Behaviour. Published by Elsevier Ltd. All rights reserved.

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Background Figs and fig-pollinating wasp species usually display a highly specific one-to-one association. However, more and more studies have revealed that the "one-to-one" rule has been broken. Co-pollinators have been reported, but we do not yet know how they evolve. They may evolve from insect speciation induced or facilitated by Wolbachia which can manipulate host reproduction and induce reproductive isolation. In addition, Wolbachia can affect host mitochondrial DNA evolution, because of the linkage between Wolbachia and associated mitochondrial haplotypes, and thus confound host phylogeny based on mtDNA. Previous research has shown that fig wasps have the highest incidence of Wolbachia infection in all insect taxa, and Wolbachia may have great influence on fig wasp biology. Therefore, we look forward to understanding the influence of Wolbachia on mitochondrial DNA evolution and speciation in fig wasps. Results We surveyed 76 pollinator wasp specimens from nine Ficus microcarpa trees each growing at a different location in Hainan and Fujian Provinces, China. We found that all wasps were morphologically identified as Eupristina verticillata, but diverged into three clades with 4.22-5.28% mtDNA divergence and 2.29-20.72% nuclear gene divergence. We also found very strong concordance between E. verticillata clades and Wolbachia infection status, and the predicted effects of Wolbachia on both mtDNA diversity and evolution by decreasing mitochondrial haplotypes. Conclusions Our study reveals that the pollinating wasp E. verticillata on F. microcarpa has diverged into three cryptic species, and Wolbachia may have a role in this divergence. The results also indicate that Wolbachia strains infecting E. verticillata have likely resulted in selective sweeps on host mitochondrial DNA.

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Lifetime reproductive success in female insects is often egg- or time-limited. For instance in pro-ovigenic species, when oviposition sites are abundant, females may quickly become devoid of eggs. Conversely, in the absence of suitable oviposition sites, females may die before laying all of their eggs. In pollinating fig wasps (Hymenoptera: Agaonidae), each species has an obligate mutualism with its host fig tree species [Ficus spp. (Moraceae)]. These pro-ovigenic wasps oviposit in individual ovaries within the inflorescences of monoecious Ficus (syconia, or ‘figs’), which contain many flowers. Each female flower can thus become a seed or be converted into a wasp gall. The mystery is that the wasps never oviposit in all fig ovaries, even when a fig contains enough wasp females with enough eggs to do so. The failure of all wasps to translate all of their eggs into offspring clearly contributes to mutualism persistence, but the underlying causal mechanisms are unclear. We found in an undescribed Brazilian Pegoscapus wasp population that the lifetime reproductive success of lone foundresses was relatively unaffected by constraints on oviposition. The number of offspring produced by lone foundresses experimentally introduced into receptive figs was generally lower than the numbers of eggs carried, despite the fact that the wasps were able to lay all or most of their eggs. Because we excluded any effects of intraspecific competitors and parasitic non-pollinating wasps, our data suggest that some pollinators produce few offspring because some of their eggs or larvae are unviable or are victims of plant defences.

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It is thought that speciation in phytophagous insects is often due to colonization of novel host plants, because radiations of plant and insect lineages are typically asynchronous. Recent phylogenetic comparisons have supported this model of diversification for both insect herbivores and specialized pollinators. An exceptional case where contemporaneous plant insect diversification might be expected is the obligate mutualism between fig trees (Ficus species, Moraceae) and their pollinating wasps (Agaonidae, Hymenoptera). The ubiquity and ecological significance of this mutualism in tropical and subtropical ecosystems has long intrigued biologists, but the systematic challenge posed by >750 interacting species pairs has hindered progress toward understanding its evolutionary history. In particular, taxon sampling and analytical tools have been insufficient for large-scale co-phylogenetic analyses. Here, we sampled nearly 200 interacting pairs of fig and wasp species from across the globe. Two supermatrices were assembled: on average, wasps had sequences from 77% of six genes (5.6kb), figs had sequences from 60% of five genes (5.5 kb), and overall 850 new DNA sequences were generated for this study. We also developed a new analytical tool, Jane 2, for event-based phylogenetic reconciliation analysis of very large data sets. Separate Bayesian phylogenetic analyses for figs and fig wasps under relaxed molecular clock assumptions indicate Cretaceous diversification of crown groups and contemporaneous divergence for nearly half of all fig and pollinator lineages. Event-based co-phylogenetic analyses further support the co-diversification hypothesis. Biogeographic analyses indicate that the presentday distribution of fig and pollinator lineages is consistent with an Eurasian origin and subsequent dispersal, rather than with Gondwanan vicariance. Overall, our findings indicate that the fig-pollinator mutualism represents an extreme case among plant-insect interactions of coordinated dispersal and long-term co-diversification.

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We report evidence that helps resolve two competing explanations for stability in the mutualism between Ficus racemosa fig trees and the Ceratosolen fusciceps wasps that pollinate them. The wasps lay eggs in the tree's ovules, with each wasp larva developing at the expense of a fig seed. Upon maturity, the female wasps collect pollen and disperse to a new tree, continuing the cycle. Fig fitness is increased by producing both seeds and female wasps, whereas short-term wasp fitness increases only with more wasps, thereby resulting in a conflict of interests. We show experimentally that wasps exploit the inner layers of ovules first (the biased oviposition explanation), which is consistent with optimal-foraging theory. As oviposition increases, seeds in the middle layer are replaced on a one-to-one basis by pollinator offspring, which is also consistent with biased oviposition. Finally, in the outer layer of ovules, seeds disappear but are only partially replaced by pollinator offspring, which suggests high wasp mortality (the biased survival or ‘unbeatable seeds’ explanation). Our results therefore suggest that both biased oviposition and biased survival ensure seed production, thereby stabilizing the mutualism. We further argue that biased oviposition can maintain biased survival by selecting against wasp traits to overcome fig defenses. Finally, we report evidence suggesting that F. racemosa balances seed and wasp production at the level of the tree. Because figs are probably selected to allocate equally to male and female function, a 1:1 seed:wasp ratio suggests that fig trees are in control of the mutualism.

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Summary 1. A trophic cascade occurs when predators directly decrease the densities, or change the behaviour, of herbivores and thus indirectly increase plant productivity. The predator–herbivore– plant context is well known, but some predators attack species beneficial to plants (e.g. pollinators) and/or enemies of herbivores (e.g. parasites), and their role in the dynamics of mutualisms remains largely unexplored. 2. We surveyed the predatory ant species and studied predation by the dominant ant species, the weaver ant Oecophylla smaragdina, associated with the fig tree Ficus racemosa in southwest China. We then tested the effects of weaver ants on the oviposition behaviour of pollinating and non-pollinating fig wasps in an ant-exclusion experiment. The effects of weaver ants on fig wasp community structure and fig seed production were then compared between trees with and without O. smaragdina. 3. Oecophylla smaragdina captured more non-pollinating wasps (Platyneura mayri) than pollinators as the insects arrived to lay eggs. When ants were excluded, more non-pollinators laid eggs into figs and fewer pollinators entered figs. Furthermore, trees with O. smaragdina produced more pollinator offspring and fewer non-pollinator offspring, shifting the community structure significantly. In addition, F. racemosa produced significantly more seeds on trees inhabited by weaver ants. 4. Oecophylla smaragdina predation reverses the dominance of the two commonest wasp species at the egg-laying stage and favours the pollinators. This behavioural pattern is mirrored by wasp offspring production, with pollinators’ offspring dominating figs produced by trees inhabited by weaver ants, and offspring of the non-pollinator P. mayri most abundant in figs on trees inhabited by other ants. 5. Overall, our results suggest that predation by weaver ants limits the success of the non-pollinating P. mayri and therefore indirectly benefits the mutualism by increasing the reproductive success of both the pollinators and the plant. Predation is thus a key functional factor that can shape the community structure of a pollinator-plant mutualistic system. Key-words: competitive release, fig wasp, mutualism, predation, predator-exclusion experiment, trophic cascade

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Endophytic insects and their parasitoids provide valuable models for community ecology. The wasp communities in inflorescences of fig trees have great potential for comparative studies, but we must first describe individual communities. Here, we add to the few detailed studies of such communities by describing the one associated with Ficus rubiginosa in Australia. First, we describe community composition, using two different sampling procedures. Overall, we identified 14 species of non-pollinating fig wasp (NPFW) that fall into two size classes. Small wasps, including pollinators, gallers and their parasitoids, were more abundant than large wasps (both galler and parasitoid species). We show that in figs where wasps emerge naturally, the presence of large wasps may partly explain the low emergence of small wasps. During fig development, large gallers oviposit first, before and around the time of pollination, while parasitoids lay eggs after pollination. We further show that parasitoids in the subfamily Sycoryctinae, which comprise the majority of all individual NPFWs, segregate temporally by laying eggs at different stages of fig development. We discuss our results in terms of species co-existence and community structure and compare our findings to those from fig wasp communities on other continents.

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We characterised a set of nine polymorphic microsatellite loci for Pleistodontes imperialis sp. 1, the pollinator wasp of Port Jackson fig (Ficus rubiginosa) in south-eastern Australia. Characterisation was performed on 30 female individuals collected from a population in Sydney, Australia. The average number of alleles per locus was 7.33, and eight loci were not in Hardy–Weinberg equilibrium. This was expected as fig wasps are known to be highly inbred. A test of genetic differentiation between two natural populations of P. imperialis sp. 1 (Sydney and Newcastle, Australia – some 120 km apart) yielded a very low FST value of 0.012, suggesting considerable gene flow. Bayesian clustering analysis using TESS 2.3.1, which does not assume Hardy–Weinberg equilibrium, however, indicated potential spatial substructuring between the Sydney and Newcastle populations, as well as within the Sydney population. The described loci were also characterised for two other species in the P. imperialis complex: P. imperialis sp. 2 (Townsville, Australia) and P. imperialis sp. 4 (Brisbane, Australia). Seven and six of the nine loci were polymorphic for P. imperialis sp. 2 and P.imperialis sp. 4, respectively.

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1. In many fig wasp species, armoured wingless males regularly engage in lethal fights for access to females inside figs, which act as discrete mating patches. 2. Kin selection generally opposes killing brothers, because their reproductive success provides indirect genetic benefits (inclusive fitness). However, siblicide may be avoided if (i) brothers do not occur in the same figs, or (ii) males avoid fighting brothers in the same fig. Alternatively, (iii) siblicide may occur because intense mate competition between brothers at the local scale overcomes kin selection effects, or (iv) males do not recognise kin. 3. A fig may also contain wasps from other closely related species and it is not known if males also fight with these individuals. 4. Nine microsatellite loci were used in the first genetic analysis of fighting in fig wasps. We assigned species and sibling identities to males and tested alternative fighting scenarios for three Sycoscapter wasp species in figs of Ficus rubiginosa. 5. Approximately 60% of figs contained males frommore than one Sycoscapter species and approximately 80% of fights were between conspecifics, but a surprising 20% were between heterospecific males. 6.Within species, fewfigs contained brothers, suggesting that females typically lay one son per fig. Overall, most males do not compete with brothers and all fights observed were between unrelated males. Key words:Competition, fighting, genetics, kin selection, microsatellites, relatedness.

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Esta dissertação teve como objetivo geral ampliar o conhecimento sobre a ecologia vegetal das matas de Restinga arenosa em substratos bem drenados no Rio Grande do Sul. Para tanto, foram realizados o estudo florístico e fitossociológico do componente arbóreo de cinco capões de Restinga e a verificação de padrões de interações mutualísticas entre aves frugívoras e as árvores, além do estudo do componente de regeneração e suas relações com o estrato arbóreo adulto. Para a amostragem da vegetação, foi usado o método de parcelas, incluindo-se todas as árvores com DAP ≥ 5cm, totalizando uma área de 1,02ha. Com estes dados, foram estimados os parâmetros usuais em fitossociologia. Em um dos capões, foi realizado também o levantamento florístico e fitossociológico das plântulas (0,05 ≤ altura < 1m) e juvenis (altura ≥ 1m, DAP < 5cm), avaliando-se as relações com o estrato arbóreo adulto, o potencial e a taxa de regeneração natural para cada espécie. Para o estudo dos mutualismos, foram feitas observações visuais e capturas de aves durante um ano. Foram estimadas a conectância do sistema mutualístico e o índice de importância das espécies. Também foi feita a rede de interações do sistema e feita a análise da variação destas interações ao longo das estações do ano. A composição florística resultou em uma riqueza total de 20 famílias e 29 espécies para os cinco capões. A densidade total arbórea teve uma média máxima de 1207 ind/ha e mínima de 747 ind/ha. Sebastiania serrata apresentou o maior valor de importância e Myrtaceae foi a família mais representada. A diversidade específica foi baixa, variando de 1,08 a 2,38 (nats). No sistema mutualístico, registraram-se 29 espécies interagindo (aves e plantas), com uma conectância de 23,9%. Turdus amaurochalinus e T. rufiventris interagiram com a maioria das espécies arbóreas e tiveram o maior índice de importância, sendo caracterizadas como as principais dispersoras em potencial. Ocotea pulchella e Myrsine spp. foram registradas com maior número de eventos de consumo de frutos, no entanto, Ficus organesis interagiu com mais espécies frugívoras, além de ter a maior importância na dieta das aves. Houve variações no número eventos de frugivoria ao longo das estações, bem como no número de espécies frugívoras e de espécies arbóreas consumidas. O componente de regeneração apresentou riqueza específica e diversidade semelhantes às do estrato arbóreo adulto, refletindo uma similaridade florística maior que 70%. A maioria das espécies (73,7%) apresentou taxa de regeneração negativa, revelando o padrão de 'J' invertido. Os resultados indicam a existência de diferenças na composição e estrutura arbórea entre os capões de Restinga, além de uma boa capacidade de regeneração para a maioria das espécies vegetais estudadas. Os dados revelam também um sistema dispersão generalista, no qual poucas espécies de aves interagem com muitas espécies arbóreas e vice-versa.

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Embora a assembléia de pequenos mamíferos desempenhe diferentes e importantes papéis na dinâmica das florestas, o conhecimento sobre a riqueza e o papel ecológico desempenhado pelas espécies nas restingas do estado do Rio Grande do Sul é ainda limitado, tendo-se poucas informações ecológicas sobre quem são esses animais nos diferentes fragmentos de florestas e como é sua relação espacial em ambientes de borda. Apesar da relevância, poucos estudos buscaram verificar se a assembléia de pequenos mamíferos em fragmentos florestais é sensível aos efeitos de borda, qual a extensão da borda para esses animais e se ela varia em diferentes estações do ano. No extremo norte da planície costeira do estado do Rio Grande do Sul, em um fragmento de floresta paludosa, denominada “Faxinal”, o presente trabalho foi realizado com o objetivo de investigar qual a riqueza e sucesso de captura de pequenos mamíferos, como é sua relação com a distância da beira do fragmento e se essa relação varia entre as estações do ano. Nos meses amostrais de janeiro (verão), abril (outono) e agosto (inverno) de 2004 foram colocadas armadilhas de arame de dois tamanhos nos estratos terrestre e arbustivo, em um gradeado com seis transecções paralelas que cobriram parte de formação florestal (3,17ha) e parte de campestre (0,31ha). A distância dos postos de captura do gradeado à beira mais próxima da floresta foram agrupadas em seis classes (1: 0-30m; 2: 31-70m; 3: 71-100; 4: 101-120; 5: 121-131; 6: 131-160m, no solo e 1: 0-20m; 2: 40-71m; 3: 80-100m; 4: 101-120m; 5: 121-131m; 6: 132-160m, no estrato arbustivo) para verificar se os dados de sucesso de captura das espécies de pequenos mamíferos diferem entre os fatores distância da beira e períodos amostrais (estações do ano). Onze espécies de pequenos mamíferos foram capturadas somente na parte florestada do gradeado (sucesso de captura total de 15,49%), sendo que maior riqueza, número de indivíduos e de capturas foram de roedores e no estrato terrestre. As espécies mais capturadas foram Akodon montensis (154 indivíduos), Oligoryzomys nigripes (34 indivíduos), Brucepattersonius iheringi (19 indivíduos), Oryzomys angouya (12 indivíduos) e Micoureus cf. demerarae (8 indivíduos) e Didelphis albiventris (7 indivíduos). O mês amostral de inverno foi aquele que apresentou maior riqueza, maior sucesso de captura e maior biomassa. As análises demonstraram relação significativa entre o sucesso de captura do conjunto das espécies de pequenos mamíferos capturadas no solo e a distância da beira, cuja relação esteve fortemente associada à estação do ano. A estrutura e/ou composição da assembléia diferiram entre as duas primeiras classes de distância, que por sua vez diferiram das demais. Isso indica que a assembléia de pequenos mamíferos, capturada no solo, é sensível à beira da floresta e que a sensibilidade varia conforme a estação do ano. Por outro lado, quando avaliada individualmente a relação para a espécie dominante Akodon montensis, o sucesso de captura do roedor não evidenciou com nitidez diferença entre as classes de distância, mas houve diferença entre os períodos amostrais, sendo o mês amostral com menor média de temperatura anual aquele com maior número de indivíduos e de capturas. Além disso, por Akodon montensis ter sido a espécie mais comum e abundante em todos períodos amostrais e em todos os pontos do gradeado, coletou-se 249 amostras fecais para investigar a capacidade dessa espécie em realizar dispersão endozoocórica As amostras coletadas foram analisadas sob lupa para verificar se existiam sementes intactas e, quando colocadas em experimento de germinação, se seriam capazes de germinar. O registro de sementes inteiras nas fezes de Akodon montensis e o tratamento dado às sementes, de não comprometer a capacidade de germinação, permite que a espécie seja, ao menos em parte, considerada legítima e efetiva dispersora de Ficus organensis e de Piper cf. solmisianum

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A headspace solid-phase microextraction (HS-SPME) procedure based on five commercialised fibres (85 μm polyacrylate – PA, 100 μm polydimethylsiloxane – PDMS, 65 μm polydimethylsiloxane/divinylbenzene – PDMS/DVB, 70 μm carbowax/divinylbenzene – CW/DVB and 85 μm carboxen/polydimethylsiloxane – CAR/PDMS) is presented for the characterization of the volatile metabolite profile of four selected Madeira island fruit species, lemon (Citrus limon), kiwi (Actinidia deliciosa), papaya (Carica papaya L.) and Chickasaw plum (Prunus angustifolia). The isolation of metabolites was followed by thermal desorption gas chromatography–quadrupole mass spectrometry (GC–qMS) methodology. The performance of the target fibres was evaluated and compared. The SPME fibre coated with CW/DVB afforded the highest extraction efficiency in kiwi and papaya pulps, while in lemon and plum the same was achieved with PMDS/DVB fibre. This procedure allowed for the identification of 80 compounds, 41 in kiwi, 24 in plums, 23 in papaya and 20 in lemon. Considering the best extraction conditions, the most abundant volatiles identified in kiwi were the intense aldehydes and ethyl esters such as (E)-2-hexenal and ethyl butyrate, while in Chicasaw plum predominate 2-hexenal, 2-methyl-4-pentenal, hexanal, (Z)-3-hexenol and cyclohexylene oxide. The major compounds identified in the papaya pulp were benzyl isothiocyanate, linalool oxide, furfural, hydroxypropanone, linalool and acetic acid. Finally, lemon was shown to be the most divergent of the four fruits, being its aroma profile composed almost exclusively by terpens, namely limonene, γ-terpinene, o-cymene and α-terpinolene. Thirty two volatiles were identified for the first time in the fruit or close related species analysed and 14 volatiles are reported as novel volatile metabolites in fruits. This includes 5 new compounds in kiwi (2-cyclohexene-1,4-dione, furyl hydroxymethyl ketone, 4-hydroxydihydro-2(3H)-furanone, 5-acetoxymethyl-2-furaldehyde and ethanedioic acid), 4 in plum (4-hydroxydihydro-2(3H)-furanone, 5-methyl-2-pyrazinylmethanol, cyclohexylene oxide and 1-methylcyclohexene), 4 in papaya (octaethyleneglycol, 1,2-cyclopentanedione, 3-methyl-1,2-cyclopentanedione and 2-furyl methyl ketone) and 2 in lemon (geranyl farnesate and safranal). It is noteworthy that among the 15 volatile metabolites identified in papaya, 3-methyl-1,2-cyclopentanedione was previously described as a novel PPARγ (peroxisome proliferator-activated receptor γ) agonist, having a potential to minimize inflammation.

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Este trabalho foi realizado para avaliar o desenvolvimento e reprodução de Tetranychus urticae em cultivares comerciais de mamão Carica papaya. Foram utilizadas cultivares do grupo Formosa (Tainung 01 e Calimosa) e do grupo Solo (Golden e Sunrise). Para iniciar o bioensaio, foi transferida uma fêmea fertilizada por disco de folha (n=50 repetições) e retirado após um período de 12h, sendo avaliada a cada 12h, registrando-se o período de incubação, duração do estágio de imaturo, longevidade e fecundidade dos adultos e viabilidade desses estágios. Os resultados indicaram que as cultivares de mamão Tainung 01, Calimosa, Sunrise e Golden são bons hospedeiros para T. urticae. O parâmetro viabilidade não sofreu influência das cultivares em todas as fases de desenvolvimento avaliadas, apresentando valores superiores a 90%. Não houve diferença estatística entre as cultivares nos parâmetros: período de pre-oviposição e viabilidade dos ovos. A cultivar Tainung 01 apresentou menor potencial hospedeiro, embora houvesse menor duração nos estágios de ovo, larva, protoninfa e ovo-adulto. Nos parâmetros de tabela de vida e fertilidade, apresentou menores valores de Ro, r m e λ e maior valor de Td. Entre as cultivares, o Sunrise apresentou um elevado potencial hospedeiro para T. urticae, pois essa cultivar proporcionou a maior produção de ovos por fêmea, maior longevidade das fêmeas, bem como a maior taxa de reprodução (maior Ro, r m e λ e menor valor de Td).