Seasonal dynamics of a drosophilid (Diptera) assemblage and its potencial as bioindicator in open environments

Drosophila Fallen, 1823 (Diptera, Drosophilidae) is for long a well-established model organism for genetics and evolutionary research. The ecology of these flies, however, has only recently been better studied. Recent papers show that Drosophila assemblies can be used as bioindicators of forested environment degradation. In this work the bioindicator potential of drosophilids was evaluated in a naturally opened environment, a coastal strand-forest (restinga). Data from nine consecutive seasonal collections revealed strong temporal fluctuation pattern of the majority of Drosophila species groups. Drosophila willistoni group was more abundant at autumns, whereas D. cardini and D. tripunctata groups were, respectively, expressive at winters and springs, and D. repleta group at both seasons. The exotic species D. simulans Sturtevant, 1919 (from D. melanogaster group) and Zaprionus indianus Gupta, 1970 were most abundant at summers. Overall, the assemblage structure did not show the same characteristics of forested or urban environments, but was similar to the forests at winters and to cities at summers. This raises the question that this locality may already been under urbanization impact. Also, this can be interpreted as an easily invaded site for exotic species, what might lead to biotic homogenization and therefore can put in check the usage of drosophilid assemblages as bioindicators at open environments.

Seasonal dynamics of a drosophilid (Diptera) assemblage and its potencial as bioindicator in open environments

Drosophila Fallen, 1823 (Diptera, Drosophilidae) is for long a well-established model organism for genetics and evolutionary research. The ecology of these flies, however, has only recently been better studied. Recent papers show that Drosophila assemblies can be used as bioindicators of forested environment degradation. In this work the bioindicator potential of drosophilids was evaluated in a naturally opened environment, a coastal strand-forest (restinga). Data from nine consecutive seasonal collections revealed strong temporal fluctuation pattern of the majority of Drosophila species groups. Drosophila willistoni group was more abundant at autumns, whereas D. cardini and D. tripunctata groups were, respectively, expressive at winters and springs, and D. repleta group at both seasons. The exotic species D. simulans Sturtevant, 1919 (from D. melanogaster group) and Zaprionus indianus Gupta, 1970 were most abundant at summers. Overall, the assemblage structure did not show the same characteristics of forested or urban environments, but was similar to the forests at winters and to cities at summers. This raises the question that this locality may already been under urbanization impact. Also, this can be interpreted as an easily invaded site for exotic species, what might lead to biotic homogenization and therefore can put in check the usage of drosophilid assemblages as bioindicators at open environments.

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.

Drosophilid collection in the Font Groga site, Barcelona (Spain)

During October 23rd and 24th and November 2012 we collected a sample of drosophilids at Font Groga (Barcelona). This site is located on the foothills of the Tibidabo mountain, which is located on the northwest edge of Barcelona and at approximately 400m above sea level. The vegetation is typical for the area, and it is mainly composed of a sparse pine forest (Pinus pinea) with some oaks (Quercus ilex) and Mediterranean brushwood. Flies were netted over 12 baits containing fermenting bananas. A large proportion of D. simulans males was found. The invasive species D. suzukii (Calabria et al. 2010; Cini et al. 2012) was detected in a non-negligible quantity. Taking into account the number of males and females, the estimated Ne for D. suzukii in the Font Groga sample was 33.70. A similar value was obtained for D. subobscura (34.97). Finally, in the study of species diversity the values obtained for H" (Shannon diversity index) and J (Shannon uniformity index) were 0.678 and 0.421, respectively. These estimates are very similar to those obtained in September 2009 in Montpellier by Calabria (2012), who reported H" = 0.679 and J = 0.422, but differ from those reported by the same author in a Font Groga sample of October 2007 (H" = 0.904 and J = 0.505).

DNA barcode discovers two cryptic species and two geographical radiations in the invasive drosophilid Zaprionus indianus

Comparing introduced to ancestral populations within a phylogeographical context is crucial in any study aiming to understand the ecological genetics of an invasive species. Zaprionus indianus is a cosmopolitan drosophilid that has recently succeeded to expand its geographical range upon three continents (Africa, Asia and the Americas). We studied the distribution of mitochondrial DNA (mtDNA) haplotypes for two genes (CO-I and CO-II) among 23 geographical populations. mtDNA revealed the presence of two well-supported phylogenetic lineages (phylads), with bootstrap value of 100%. Phylad I included three African populations, reinforcing the African-origin hypothesis of the species. Within phylad II, a distinct phylogeographical pattern was discovered: Atlantic populations (from the Americas and Madeira) were closer to the ancestral African populations than to Eastern ones (from Madagascar, Middle East and India). This means that during its passage from endemism to cosmopolitanism, Z. indianus exhibited two independent radiations, the older (the Eastern) to the East, and the younger (the Atlantic) to the West. Discriminant function analysis using 13 morphometrical characters was also able to discriminate between the two molecular phylads (93.34 +/- 1.67%), although detailed morphological analysis of male genitalia using scanning electron microscopy showed no significant differences. Finally, crossing experiments revealed the presence of reproductive barrier between populations from the two phylads, and further between populations within phylad I. Hence, a bona species status was assigned to two new, cryptic species: Zaprionus africanus and Zaprionus gabonicus, and both were encompassed along with Z. indianus and Zaprionus megalorchis into the indianus complex. The ecology of these two species reveals that they are forest dwellers, which explains their restricted endemic distribution, in contrast to their relative cosmopolitan Z. indianus, known to be a human-commensal. Our results reconfirm the great utility of mtDNA at both inter- and intraspecific analyses within the frame of an integrated taxonomical project.

Molecular cloning of a family of xenobiotic-inducible drosophilid cytochrome P450s: Evidence for involvement in host-plant allelochemical resistance

Cytochrome P450s constitute a superfamily of genes encoding mostly microsomal hemoproteins that play a dominant role in the metabolism of a wide variety of both endogenous and foreign compounds. In insects, xenobiotic metabolism (i.e., metabolism of insecticides and toxic natural plant compounds) is known to involve members of the CYP6 family of cytochrome P450s. Use of a 3′ RACE (rapid amplification of cDNA ends) strategy with a degenerate primer based on the conserved cytochrome P450 heme-binding decapeptide loop resulted in the amplification of four cDNA sequences representing another family of cytochrome P450 genes (CYP28) from two species of isoquinoline alkaloid-resistant Drosophila and the cosmopolitan species Drosophila hydei. The CYP28 family forms a monophyletic clade with strong regional homologies to the vertebrate CYP3 family and the insect CYP6 family (both of which are involved in xenobiotic metabolism) and to the insect CYP9 family (of unknown function). Induction of mRNA levels for three of the CYP28 cytochrome P450s by toxic host-plant allelochemicals (up to 11.5-fold) and phenobarbital (up to 49-fold) corroborates previous in vitro metabolism studies and suggests a potentially important role for the CYP28 family in determining patterns of insect–host-plant relationships through xenobiotic detoxification.

Variations on a theme: diversification of cuticular hydrocarbons in a clade of cactophilic Drosophila

Background: We characterized variation and chemical composition of epicuticular hydrocarbons (CHCs) in the seven species of the Drosophila buzzatii cluster with gas chromatography/mass spectrometry. Despite the critical role of CHCs in providing resistance to desiccation and involvement in communication, such as courtship behavior, mating, and aggregation, few studies have investigated how CHC profiles evolve within and between species in a phylogenetic context. We analyzed quantitative differences in CHC profiles in populations of the D. buzzatii species cluster in order to assess the concordance of CHC differentiation with species divergence. Results: Thirty-six CHC components were scored in single fly extracts with carbon chain lengths ranging from C(29) to C(39), including methyl-branched alkanes, n alkenes, and alkadienes. Multivariate analysis of variance revealed that CHC amounts were significantly different among all species and canonical discriminant function (CDF) analysis resolved all species into distinct, non-overlapping groups. Significant intraspecific variation was found in different populations of D. serido suggesting that this taxon is comprised of at least two species. We summarized CHC variation using CDF analysis and mapped the first five CHC canonical variates (CVs) onto an independently derived period (per) gene + chromosome inversion + mtDNA COI gene for each sex. We found that the COI sequences were not phylogenetically informative due to introgression between some species, so only per + inversion data were used. Positive phylogenetic signal was observed mainly for CV1 when parsimony methods and the test for serial independence (TFSI) were used. These results changed when no outgroup species were included in the analysis and phylogenetic signal was then observed for female CV3 and/or CV4 and male CV4 and CV5. Finally, removal of divergent populations of D. serido significantly increased the amount of phylogenetic signal as up to four out of five CVs then displayed positive phylogenetic signal. Conclusions: CHCs were conserved among species while quantitative differences in CHC profiles between populations and species were statistically significant. Most CHCs were species-, population-, and sex-specific. Mapping CHCs onto an independently derived phylogeny revealed that a significant portion of CHC variation was explained by species' systematic affinities indicating phylogenetic conservatism in the evolution of these hydrocarbon arrays, presumptive waterproofing compounds and courtship signals as in many other drosophilid species.

Subtropical drosophilids in Australia can be characterized by adult distribution across vegetation type and by height above forest floor

Understanding the pattern in which adult drosophilids of different species are distributed across and within different vegetation types is necessary for accurate interpretation of their local ecology and diversity. Such studies have been conducted mainly in temperate regions, and there is no basis for extrapolating their conclusions to tropical areas. This study describes the vertical distribution (0-20 m) of drosophilids attracted to banana baits in five different vegetation types in subtropical eastern Australia including open woodland, and rain-forest types. The distribution of most of the 15 common species could be characterized three-dimensionally by vegetation type and height above forest floor. Only one species, Scaptodrosophila lativittata, was common in all vegetation types and it was a canopy species in rain forests and a ground-level species in open woodland. Vertical distribution of some species clearly matched that of their larval hosts, but it did not in others. For example, the fungivore Leucophenga scutellata was mostly trapped well above the forest floor, yet it breeds at ground level, suggesting behavioural mode can influence vertical distributions. We conclude that the vertical dimension, although still poorly understood in relation to drosophilid habitats, needs to be taken into account when conducting and interpreting studies aimed at understanding drosophilid populations and communities in the subtropics.

Larval hosts of Australian Drosophilidae (Diptera): A field survey in subtropical and tropical Australia

The drosophilid fauna in Australia offers an important study system for evolutionary studies. Larval hosts are unknown for most species, however, and this imposes serious limits to understanding their ecological context. The present paper reports the first systematic, large-scale field survey of potential larval hosts to be conducted, in order to obtain an overview of the host utilisation patterns of Australian drosophilids. Potential hosts (mostly fruit and fungi) were collected from different vegetation types in northern and eastern Australia. Host data were obtained for 81 drosophilid species from 17 genera (or 28% of the known Fauna). Most genera were restricted to either fruit or fungi, although Scaptodrosophila spp. and Drosophila spp. were recorded from fruit, fungi, flowers and compost, and Drosophila spp. also emerged from the parasitic plant Balanophora fungosa. There was no evidence that use of either fruit or fungi was correlated to host phylogeny. Drosophilids emerged from hosts collected from all sampled vegetation types (rainforest, open forest, heath and domestic environments). Vegetation type influenced drosophilid diversity, both by affecting host availability and because some drosophilid species apparently restricted their search for hosts to particular vegetation types.

Wolbachia-mediated sperm modification is dependent on the host genotype in Drosophila

Estimates of Wolbachia density in the eggs, testes and whole flies of drosophilid hosts have been unable to predict the lack of cytoplasmic incompatibility (CI) expression in so-called mod(-) variants. Consequently, the working hypothesis has been that CI expression, although related to Wolbachia density, is also governed by unknown factors that are influenced by both host and bacterial genomes. Here, we compare the behaviour of the mod(-) over-replicating Wolbachia popcorn strain in its native Drosophila melanogaster host to the same strain transinfected into a novel host, namely Drosophila simulans. We report that (i) the popcorn strain is a close relative of other D. melanogaster infections, (ii) the mod(-) status of popcorn in D. melanogaster appears to result from its inability to colonize sperm bundles, (iii) popcorn is present in the bundles in D. simulans and induces strong CI expression, which demonstrates that the bacterial strain does not lack the genetic machinery for inducing CI and that there is host-species-specific control over Wolbachia tissue tropism, and (iv) infection of sperm bundles by the mod(-) D. simulans wCof strain indicates that there are several independent routes by which a strain can be a CI non-expressor.

Drosophilidae (Diptera) of Australia's Northern Territory: Ecology and biogeography

The drosophilid fauna is well documented in eastern Australia but is poorly known in other parts of the continent. This paper summarises what is known of this fauna in the Northern Territory (NT), and includes results from banana trapping in the humid and arid zones. The 42 recorded species include species that breed in fruit, fungi and/or flowers, and a larval predator of scale insects. Drosophilids occur in all three major climate zones (humid, semiarid and arid) but predominate in the humid zone. Banana-attracted species in the humid zone (wet-dry tropics) were common in all sampled habitats: urban, rainforest and open woodland. They included predominantly urban and/or rainforest species. Of the species collected in open woodland, some are likely to be breeding there, whereas others may have been intercepted during movement across the area. The semiarid fauna is a depauperate version of that found in the humid region. Only three species have been recorded in the arid region: an endemic arid specialist, and two cosmopolitan species (D. simulans and D. melanogaster ) in urban Alice Springs. Overall, the NT drosophilid fauna represents a depauperate version of that found in eastern Australia, probably because of climatic factors and natural barriers to range expansion. There is little evidence of regional endemism, with probably only one (and at most three) species endemic to the NT, and no evidence of independent, natural dispersion from nearby Indonesia.

Ecology of colonizing populations of the figfly Zaprionus indianus (Diptera, Drosophilidae) in Porto Alegre, Southern Brazil

Ecological parameters such as trophic niche overlap and niche breadth were computed from the frequency of all drosophilid species collected in 2001 and 2002, in the urban area of Porto Alegre city, Brazil. Independently of place, drosophilids community composition and components breeding or feeding sites, the highest frequencies of Zaprionus indianus (Gupta, 1970) were obtained during seasons with higher average temperatures, especially during summer. From a total of the 19,146 individuals of Z. indianus sampled, 80% were found in this season. The data of trophic niche breadth suggest a higher opportunism of adult flies in feeding substrate use, but the females are still selective in oviposition substrate choice. For the values of niche overlap, only for some species, did we find any correlation between these and species richness and/or number of resources, and the most of the correlation values considering breeding and feeding sites were positive. The colonization of Z. indianus in Porto Alegre could be leading to adjustments in the survival strategies of the resident species, but it is possible that many species have conditions for coexisting with the invader.

Two decades of colonization of the urban environment of Porto Alegre, southern Brazil, by Drosophila paulistorum (Diptera, Drosophilidae)

Drosophila paulistorum Dobzhansky & Pavan, 1949 had initially been considered absent in anthropogenically disturbed environments, but in 1985 the detection of the species in Porto Alegre city, southern Brazil, suggested its potential to colonize new habitats and laid the foundations for ecologic studies on this species' populations. This study followed the variations in D. paulistorum populations in this town almost 20 years after its first local record. Drosophilid specimens were collected in sites with different urbanization grades and the results point to the expressive decline in D. paulistorum populations in Porto Alegre. This decline may be linked to urban growth and to naturally driven population decline, as imputed to climatic changes like variations in maximum and minimum temperatures as a consequence of a global climate warming. Also, the recent introduction of exotic species Zaprionus indianus Gupta, 1970 seems to play a role in this scenario, changing the interactions between native species.

Assemblages of drosophilids (Diptera, Drosophilidae) in mangrove forests: community ecology and species diversity

Brazilian fauna of drosophilids has been researched in various ecosystems, but those in mangrove forests remain overlooked in Brazil and elsewhere. The present study attempts to characterise the assemblages of drosophilids of this environment, based on 28 collections taken in three mangrove areas in Santa Catarina Island, southern Brazil. The three mangroves surveyed were different in their surroundings, which varied from highly urbanised areas to conservation areas with natural vegetation. Overall, 69 species were collected, and no remarkable difference was detected in species composition and abundances or in the richness, evenness and heterogeneity between sites. The species abundance distribution observed fitted to a theoretical lognormal distribution in the three mangroves. The species richness scored and the performance of the species richness estimators showed an unexpectedly high diversity, considering the very low floristic diversity and the harsh conditions of the environment. Regarding species composition and abundances, the drosophilid mangrove assemblages were shown to be more similar to those found in open environments, with a marked dominance of exotic species. Finally, considering the apparent lack of feeding and breeding sites, we suggest that mangrove forests are acting as sink habitats for the drosophilids populations.

Chemosensory ecology: deceiving Drosophila.

The Solomon's lily arum mimics the odours of yeast to attract drosophilid flies as unrewarded pollinators.