A remarkable legion of guests: Diversity and host specificity of army ant symbionts.

Tropical rainforests are among the most diverse biomes on Earth. While species inventories are far from complete for any tropical rainforest, even less is known about the intricate species interactions that form the basis of these ecological communities. One fascinating but poorly studied example are the symbiotic associations between army ants and their rich assemblages of parasitic arthropod guests. Hundreds of these guests, or myrmecophiles, have been taxonomically described. However, because previous work has mainly been based on haphazard collections from disjunct populations, it remains challenging to define species boundaries. We therefore know little about the species richness, abundance and host specificity of most guests in any given population, which is crucial to understand co-evolutionary and ecological dynamics. Here, we report a quantitative community survey of myrmecophiles parasitizing the six sympatric Eciton army ant species in a Costa Rican rainforest. Combining DNA barcoding with morphological identification of over 2,000 specimens, we discovered 62 species, including 49 beetles, 11 flies, one millipede and one silverfish. At least 14 of these species were new to science. Ecological network analysis revealed a clear signal of host partitioning, and each Eciton species was host to both specialists and generalists. These varying degrees in host specificities translated into a moderate level of network specificity, highlighting the system's level of biotic pluralism in terms of biodiversity and interaction diversity. By providing vouchered DNA barcodes for army ant guest species, this study provides a baseline for future work on co-evolutionary and ecological dynamics in these species-rich host-symbiont networks across the Neotropical realm.

Phylogeny, host association and biogeographical patterns in the diverse millipede-parasitoid genus Myriophora Brown (Diptera: Phoridae).

Myriophora is the most species-rich group of parasitoids that attack toxic, chemically defended millipedes in the superorder Juliformia and order Polydesmida-a resource that few insect predators and parasitoids are able to exploit. Worldwide, there are an estimated 200 species of Myriophora, with the majority of the diversity centred in the Neotropical region. The phylogeny of Myriophora is unknown, biogeographical patterns are not documented, and known host associations have not been assessed in a phylogenetic context. We provide the first phylogenetic study of the genus from a data set composed of 52 taxa primarily from the Neotropical region including 10 outgroups, 40 morphological characters, and molecular data from three mitochondrial (16S, COI and ND1) and one nuclear marker (AK). We find that Myriophora dispersed from the New World to the Old World in a single event before subsequently spreading to the Afrotropical region. The ancestral hosts reconstructed for Myriophora are the benzoquinone-producing Juliformia, and this association has been retained in the Old World clade. In the Neotropical region, Myriophora that are associated with cyanide-producing polydesmidan millipedes are confined to a single clade that shows remarkably little genetic variation between clearly morphologically diagnosable species.

Millipede Defensive Compounds Are a Double-Edged Sword: Natural History of the Millipede-Parasitic Genus Myriophora Brown (Diptera: Phoridae).

Toxic defensive secretions produced by millipedes in the orders Julida, Spirobolida, Spirostreptida, and Polydesmida are highly repellent to most vertebrate and invertebrate natural enemies, but a few insects have evolved mechanisms to overcome these defenses. We demonstrate that highly specialized parasitic phorid flies in the species-rich genus Myriophora use volatile millipede defensive compounds as kairomones for host location. Of the two predominant quinone components in the defensive blend of juliform millipedes, 2-methoxy-3-methyl-1,4-benzoquinone alone was sufficient to attract adult flies of both sexes; however, a combination of 2-methoxy-3-methyl-1,4-benzoquinone and 2-methyl-1,4-benzoquinone increased attractiveness nearly threefold. We further discuss oviposition behavior, adult and larval feeding habits, life history parameters, and the potential competitive interactions between phorid flies in the genus Myriophora and other millipede-associated insects.

The Attractiveness of the Gravid Aedes Trap to Dengue Vectors in Fiji.

The Gravid Aedes Trap (GAT) is a passive trap that relies on visual and olfactory cues to lure and capture gravid mosquitoes. The GAT was designed as a dengue vector surveillance tool for use in conditions where power sources are not readily available. Experiments were conducted over a 10-d period in Fiji to determine which species of mosquitoes the GAT would collect, how different infusion types affect the attractiveness of the trap, how long each infusion type took to begin attracting mosquitoes, and how long the infusion was attractive to container breeding Aedes mosquitoes. Infusions were created by adding 10 g of organic material to 2 liters of water. Infusions were made using chicken feed pellets, dried mango leaves (Mangifera indica), and dried lawn grasses (primarily Axonopus spp.). The GAT collected four different vectors Aedes aegypti (Linnaeus) (Diptera: Culicidae), Aedes albopictus (Skuse) (Diptera: Culicidae), Aedes polynesiensis Mark (Diptera: Culicidae), and Culex quinquefasciatus Say (Diptera: Culicidae). As observed in previous studies, using some type of organic infusion improved the attractiveness of the trap. Of the three different infusion components tested, chicken feed was most attractive followed by lawn grasses and mango leaves. All infusions performed better than plain water. Chicken feed was found to be most attractive for Ae. aegypti and Ae. albopictus during the first 6 d after placing the traps and for Cx. quinquifasciatus in days 7-9. The mango and grass infusions took longer to attract mosquitoes but were most attractive during days 7-10 for all species.

Comprehensive inventory of true flies (Diptera) at a tropical site.

Estimations of tropical insect diversity generally suffer from lack of known groups or faunas against which extrapolations can be made, and have seriously underestimated the diversity of some taxa. Here we report the intensive inventory of a four-hectare tropical cloud forest in Costa Rica for one year, which yielded 4332 species of Diptera, providing the first verifiable basis for diversity of a major group of insects at a single site in the tropics. In total 73 families were present, all of which were studied to the species level, providing potentially complete coverage of all families of the order likely to be present at the site. Even so, extrapolations based on our data indicate that with further sampling, the actual total for the site could be closer to 8000 species. Efforts to completely sample a site, although resource-intensive and time-consuming, are needed to better ground estimations of world biodiversity based on limited sampling.

Remarkable fly (Diptera) diversity in a patch of Costa Rican cloud forest: Why inventory is a vital science.

Study of all flies (Diptera) collected for one year from a four-hectare (150 x 266 meter) patch of cloud forest at 1,600 meters above sea level at Zurquí de Moravia, San José Province, Costa Rica (hereafter referred to as Zurquí), revealed an astounding 4,332 species. This amounts to more than half the number of named species of flies for all of Central America. Specimens were collected with two Malaise traps running continuously and with a wide array of supplementary collecting methods for three days of each month. All morphospecies from all 73 families recorded were fully curated by technicians before submission to an international team of 59 taxonomic experts for identification.        Overall, a Malaise trap on the forest edge captured 1,988 species or 51% of all collected dipteran taxa (other than of Phoridae, subsampled only from this and one other Malaise trap). A Malaise trap in the forest sampled 906 species. Of other sampling methods, the combination of four other Malaise traps and an intercept trap, aerial/hand collecting, 10 emergence traps, and four CDC light traps added the greatest number of species to our inventory. This complement of sampling methods was an effective combination for retrieving substantial numbers of species of Diptera. Comparison of select sampling methods (considering 3,487 species of non-phorid Diptera) provided further details regarding how many species were sampled by various methods.        Comparison of species numbers from each of two permanent Malaise traps from Zurquí with those of single Malaise traps at each of Tapantí and Las Alturas, 40 and 180 km distant from Zurquí respectively, suggested significant species turnover. Comparison of the greater number of species collected in all traps from Zurquí did not markedly change the degree of similarity between the three sites, although the actual number of species shared did increase.        Comparisons of the total number of named and unnamed species of Diptera from four hectares at Zurquí is equivalent to 51% of all flies named from Central America, greater than all the named fly fauna of Colombia, equivalent to 14% of named Neotropical species and equal to about 2.7% of all named Diptera worldwide. Clearly the number of species of Diptera in tropical regions has been severely underestimated and the actual number may surpass the number of species of Coleoptera.        Various published extrapolations from limited data to estimate total numbers of species of larger taxonomic categories (e.g., Hexapoda, Arthropoda, Eukaryota, etc.) are highly questionable, and certainly will remain uncertain until we have more exhaustive surveys of all and diverse taxa (like Diptera) from multiple tropical sites.        Morphological characterization of species in inventories provides identifications placed in the context of taxonomy, phylogeny, form, and ecology. DNA barcoding species is a valuable tool to estimate species numbers but used alone fails to provide a broader context for the species identified.

Baby Killers: Documentation and Evolution of Scuttle Fly (Diptera: Phoridae) Parasitism of Ant (Hymenoptera: Formicidae) Brood.

BACKGROUND: Numerous well-documented associations occur among species of scuttle flies (Diptera: Phoridae) and ants (Hymenoptera: Formicidae), but examples of brood parasitism are rare and the mechanisms of parasitism often remain unsubstantiated. NEW INFORMATION: We present two video-documented examples of ant brood (larvae and pupae) parasitism by scuttle flies. In footage from Estação Biológica de Boracéia in Brazil, adult females of Ceratoconus setipennis Borgmeier can be seen attacking workers of Linepithema humile (Mayr) species group while they are carrying brood, and ovipositing directly onto brood in the nest. In another remarkable example, footage from the Soltis Center, near Peñas Blancas in Costa Rica, shows adult females of an unidentified species of the Apocephalus grandipalpus Borgmeier group mounting Pheidole Westwood brood upside-down and ovipositing while the brood are being transported by workers. Analysis of evolutionary relationships (in preparation) among Apocephalus Coquillett species shows that this is a newly derived behavior within the genus, as the A. grandipalpus group arises within a group of adult ant parasitoids. In contrast, relationships of Ceratoconus Borgmeier have not been studied, and the lifestyles of the other species in the genus are largely unknown.

Revision of the New World Species of the Millipede-Parasitic Genus Myriophora Brown (Diptera: Phoridae).

The New World species of the millipede-parasitic genus Myriophora Brown are revised. Sixty-five species based on the female sex are treated, mostly from the Neotropical Region. Of these, fifty-seven are new to science: Myriophora alexandrae sp. nov., Myriophora alienipennis sp. nov., Myriophora angustifascia sp. nov., Myriophora annetteae sp. nov., Myriophora annulata sp. nov., Myriophora bicuspidis sp. nov., Myriophora bilsae sp. nov., Myriophora bimaculata sp. nov., Myriophora borealis sp. nov., Myriophora brevitarsus sp. nov., Myriophora browni sp. nov. Hash, Myriophora brunneipleuron sp. nov., Myriophora communis sp. nov., Myriophora curvata sp. nov., Myriophora curvicacumen sp. nov., Myriophora dennisoni sp. nov., Myriophora discalis sp. nov., Myriophora diversa sp. nov., Myriophora dividida sp. nov., Myriophora dolionatis sp. nov., Myriophora flavicosta sp. nov., Myriophora fuscidorsum sp. nov., Myriophora gigantea sp. nov., Myriophora gobaleti sp. nov., Myriophora harwoodi sp. nov., Myriophora hebes sp. nov., Myriophora heratyi sp. nov., Myriophora inaequalisetarum sp. nov., Myriophora infirmata sp. nov., Myriophora jeffersoni sp. nov., Myriophora kerri sp. nov., Myriophora kungae sp. nov., Myriophora longisetarum sp. nov., Myriophora luteitergum sp. nov., Myriophora magnilabellum sp. nov., Myriophora misionesensis sp. nov., Myriophora nigra sp. nov., Myriophora nigralinea sp. nov., Myriophora obscuritergum sp. nov., Myriophora pallida sp. nov., Myriophora parva sp. nov., Myriophora pectinata sp. nov., Myriophora perpendicularis sp. nov., Myriophora plana sp. nov., Myriophora porrasae sp. nov., Myriophora porrecta sp. nov., Myriophora reminatis sp. nov., Myriophora scopulata sp. nov., Myriophora simplex sp. nov., Myriophora sinesplendida sp. nov., Myriophora smithi sp. nov., Myriophora spicaphora sp. nov., Myriophora spicaticonus sp. nov., Myriophora tenuis sp. nov., Myriophora uruguaiensis sp. nov., Myriophora vancouverensis sp. nov., Myriophora wellsorum sp. nov. The first key to the species of Myriophora is provided.