Ratio changes in blood-feeding and voltinism in black flies (Diptera: Simuliidae) over a latitudinal gradient in North America north of Mexico.

We explored how the ratio of blood-feeding behaviour-nonfeeding, bird feeding (ornithophily) and mammal feeding (mammalophily)-and voltinism (univoltine and multivoltine) vary over a latitudinal gradient from Alaska to Florida. These two fixed species traits were divided into five mutually exclusive combinations of trait states-nonfeeding/univoltine, ornithophilic/univoltine, mammalophilic/univoltine, ornithophilic/multivoltine and mammalophilic/multivoltine-within each of three datasets (north, east and west). We found a significant association between location (north, east and west) and trait state, which was driven by the large percentage of nonfeeding females in the north. When this trait state was removed, no significant differences were found for the remaining trait states and locations. Although the distribution of trait states did not differ between east and west datasets, the distribution with relation to 1° changes in latitude within each of these datasets showed distinct differences. In the east, both ornithophilic/univoltine and mammalophilic/univoltine species significantly increased with latitude, in proportion to the total species present. In contrast, the proportion of mammalophilic/multivoltine species decreased as latitude increased. Ornithophilic/multivoltine species in the east and the trait states in the west did not show any significant relationship to latitude.

Haemolymph viscosity in hawkmoths and its implications for hovering flight.

Viscosity determines the resistance of haemolymph flow through the insect body. For flying insects, viscosity is a major physiological parameter limiting flight performance by controlling the flow rate of fuel to the flight muscles, circulating nutrients and rapidly removing metabolic waste products. The more viscous the haemolymph, the greater the metabolic energy needed to pump it through confined spaces. By employing magnetic rotational spectroscopy with nickel nanorods, we showed that viscosity of haemolymph in resting hawkmoths (Sphingidae) depends on wing size non-monotonically. Viscosity increases for small hawkmoths with high wingbeat frequencies, reaches a maximum for middle-sized hawkmoths with moderate wingbeat frequencies, and decreases in large hawkmoths with slower wingbeat frequencies but greater lift. Accordingly, hawkmoths with small and large wings have viscosities approaching that of water, whereas hawkmoths with mid-sized wings have more than twofold greater viscosity. The metabolic demands of flight correlate with significant changes in circulatory strategies via modulation of haemolymph viscosity. Thus, the evolution of hovering flight would require fine-tuned viscosity adjustments to balance the need for the haemolymph to carry more fuel to the flight muscles while decreasing the viscous dissipation associated with its circulation.

Wettability and morphology of proboscises interweave with hawkmoth evolutionary history.

Hovering hawkmoths expend significant energy while feeding, which should select for greater feeding efficiency. Although increased feeding efficiency has been implicitly assumed, it has never been assessed. We hypothesized that hawkmoths have proboscises specialized for gathering nectar passively. Using contact angle and capillary pressure to evaluate capillary action of the proboscis, we conducted a comparative analysis of wetting and absorption properties for 13 species of hawkmoths. We showed that all 13 species have a hydrophilic proboscis. In contradistinction, the proboscises of all other tested lepidopteran species have a wetting dichotomy with only the distal ∼10% hydrophilic. Longer proboscises are more wettable, suggesting that species of hawkmoths with long proboscises are more efficient at acquiring nectar by the proboscis surface than are species with shorter proboscises. All hawkmoth species also show strong capillary pressure, which, together with the feeding behaviors we observed, ensures that nectar will be delivered to the food canal efficiently. The patterns we found suggest that different subfamilies of hawkmoths use different feeding strategies. Our comparative approach reveals that hawkmoths are unique among Lepidoptera and highlights the importance of considering the physical characteristics of the proboscis to understand the evolution and diversification of hawkmoths.

Lepidopteran mouthpart architecture suggests a new mechanism of fluid uptake by insects with long proboscises.

Proboscises of many fluid-feeding insects share a common architecture: they have a partially open food canal along their length. This feature has never been discussed in relation to the feeding mechanism. We formulated and solved a fluid mechanics model of fluid uptake and estimated the time required to completely fill the food canal of the entire proboscis through the openings along its length. Butterflies and moths are taken as illustrative and representative of fluid-feeding insects. We demonstrated that the proposed mechanism of filling the proboscis with fluid through permeable lengthwise bands, in association with a thin film of saliva in the food canal, offers a competitive pathway for fluid uptake. Compared with the conventional mechanism of fluid uptake through apically restricted openings, the new mechanism provides a faster rate of fluid uptake, especially for long-tongued insects. Accordingly, long-tongued insects with permeable lengthwise bands would be able to more rapidly exploit a broader range of liquids in the form of films, pools, and discontinuous columns, thereby conserving energy and minimizing exposure to predators, particularly for hovering insects.

The general architecture of black fly-parasite interactions: Parasitism in lotic systems at a continental scale.

We examined the general architecture of interactions between stream-dwelling larval black flies (Diptera: Simuliidae) and their common parasites in 1736 collections across North America. Mermithid nematodes (family Mermithidae), microsporidia (phylum Microsporidia), and the fungus Coelomycidium simulii Debaisieux (phylum Blastocladiomycota) infected larval black flies. We found similar continental distributions for these three parasite taxa across North America. At least one of these taxa was represented in 42.2% of all black fly collections. Species interactions in ecological networks typically imply that each link between species is equally important. By employing quantitative measures of host susceptibilities and parasite dependencies, we provide a more complete structure for host-parasite networks. The distribution of parasite dependencies and host susceptibilities were right-skewed, with low values indicating that most dependencies (parasites) and susceptibilities (hosts) were weak. Although regression analysis between host frequency and parasite incidence were highly significant, frequency analysis suggested that the distributions of parasites differ significantly among the four most common and closely related (same subgenus) species of hosts. A highly significant pattern of nestedness in our bipartite host-parasite network indicated that specialized parasites (i.e., those that interact with few host species) tend to occur as subsets of the most common hosts.

Sexually anomalous individuals of the black fly Simulium trangense (Diptera: Simuliidae) infected with mermithid parasites (Nematoda: Mermithidae).

Sexually anomalous individuals, typically intersexes or gynandromorphs, bear a mixture of male and female traits. Twelve sexually anomalous individuals of the black fly Simulium (Gomphostilbia) trangense Jitklang, Kuvangkadilok, Baimai, Takaoka & Adler were discovered among 49 adults reared from pupae. All 12 sexually anomalous adults were parasitized by mermithid nematodes, although five additional parasitized adults had no overt external anomalies. Sequence analysis of the 18S rRNA gene revealed that the mermithids, possibly representing a new species, are related to Mesomermis spp., with genetic distances of 5.09-6.87%. All 12 anomalous individuals had female phenotypical traits on the head, thorax, forelegs, midlegs, and claws, but male features on the left and right hind basitarsi. One individual had mixed male and female genitalia. The findings are in accord with the trend that mermithid infections are associated with sexually anomalous adult black flies.

The dilemma of pest suppression in the conservation of endangered species.

In the conservation of endangered species, suppression of a population of one native species to benefit another poses challenges. Examples include predator control and nest parasite reduction. Less obvious is the control of blood-feeding arthropods. We conducted a case study of the effect of native black flies (Simulium spp.) on reintroduced Whooping Cranes (Grus americana). Our intent was to provide a science-driven approach for determining the effects of blood-feeding arthropods on endangered vertebrates and identifying optimal management actions for managers faced with competing objectives. A multiyear experiment demonstrated that black flies reduce nest success in cranes by driving incubating birds off their nests. We used a decision-analytic approach to develop creative management alternatives and evaluate trade-offs among competing objectives. We identified 4 management objectives: establish a self-sustaining crane population, improve crane well-being, maintain native black flies as functional components of the ecosystem, and minimize costs. We next identified potential management alternatives: do nothing, suppress black flies, force crane renesting to occur after the activity period of black flies, relocate releases of cranes, suppress black flies and relocate releases, or force crane renesting and relocate releases. We then developed predictions on constructed scales of 0 (worst-performing alternative) to 1 (best-performing alternative) to indicate how alternative actions performed in terms of management objectives. The optimal action depended on the relative importance of each objective to a decision maker. Only relocating releases was a dominated alternative, indicating that it was not optimal regardless of the relative importance of objectives. A rational decision maker could choose any other management alternative we considered. Recognizing that decisions involve trade-offs that must be weighed by decision makers is crucial to identifying alternatives that best balance multiple management objectives. Given uncertainty about the population dynamics of blood-feeding arthropods, an adaptive management approach could offer substantial benefits.

El Dilema de la Eliminación de Plagas en la Conservación de Especies en Peligro Resumen En la conservación de especies en peligro, la eliminación de la población de una especie nativa para beneficiar a otra representa un reto. Los ejemplos incluyen el control de depredadores y la reducción de parásitos de nido. El control de artrópodos hematófagos es menos obvio. Realizamos un estudio de caso sobre el efecto que las moscas negras nativas (Simulium spp.) tienen sobre las grullas trompeteras (Grus americana) reintroducidas. Nuestra intención era proporcionar una estrategia científica para determinar los efectos que tienen los artrópodos hematófagos sobre los vertebrados en peligro de extinción e identificar las acciones de manejo óptimo para los administradores que enfrentan objetivos en competencia. Un experimento multianual demostró que las moscas negras reducen el éxito de anidación de las grullas al ahuyentar a las aves incubadoras de sus nidos. Usamos una estrategia analítica de decisión para desarrollar alternativas creativas de manejo y para evaluar las compensaciones entre los objetivos en competencia. Identificamos cuatro objetivos de manejo: establecer una población autosustentable de grullas, mejorar el bienestar de las grullas, mantener a las moscas negras nativas como componentes funcionales del ecosistema, y minimizar los costos. Después, identificamos las alternativas potenciales de manejo: no realizar acciones, eliminar a las moscas, obligar a que la re-anidación de las grullas ocurra después del periodo de actividad de las moscas, reubicar las liberaciones de las grullas, eliminar a las moscas y reubicar las liberaciones, o forzar la re-anidación de las grullas y reubicar las liberaciones. Finalmente, desarrollamos predicciones sobre escalas construidas de 0 (la alternativa con el peor desempeño) a 1 (la alternativa con el mejor desempeño) para indicar el éxito de las acciones alternativas con respecto a los objetivos de manejo. La acción óptima dependió de la importancia relativa que cada objetivo tenía para un responsable de la conservación. Sólo la reubicación de las liberaciones fue una alternativa dominada, lo que indica que no era óptima a pesar de la importancia relativa de sus objetivos. Un encargado razonable podría entonces elegir de entre cualquier otra de las alternativas de manejo que consideramos. El reconocimiento de las decisiones que involucran compensaciones que deben ser sopesadas por los encargados de la conservación es importante para la identificación de alternativas que balancean de mejor manera los objetivos múltiples del manejo. Dada la incertidumbre sobre las dinámicas poblacionales de los artrópodos hematófagos, una estrategia de manejo adaptativo podría ofrecer beneficios sustanciales.

A novel molecular and chromosomal lineage of the anthropophilic Simulium (Simulium) rufibasis subgroup (Diptera: Simuliidae) in Taiwan.

The Simulium rufibasis subgroup is one of three subgroups of the Simulium (Simulium) tuberosum species-group; it is characterized by a pair of clustered stout hairs on the ventral surface of female abdominal segment 7. A member of the S. rufibasis subgroup in Taiwan was investigated morphologically and genetically using the universal cytochrome c oxidase subunit I (COI) barcoding gene and polytene chromosomal banding pattern. The Taiwanese material is morphologically similar to S. rosliramlii Takaoka & Chen from Vietnam and represents the second species of the S. rufibasis subgroup known from Taiwan. It also represents a novel molecular lineage that is distinct from three other primary lineages identified as S. doipuiense, S. doipuiense/S. rufibasis, and S. weji previously reported from Thailand. The mitochondrial evidence for a distinct lineage in Taiwan is supported by chromosomal analysis, which revealed unique sex chromosomes. For nomenclatural stability, we associate the name S. arisanum Shiraki with the Taiwanese entity. Originally described from females from Taiwan, S. arisanum until now has remained an enigmatic species.

Estimating Diversity of Black Flies in the Simulium ignescens and Simulium tunja Complexes in Colombia: Chromosomal Rearrangements as the Core of Integrative Taxonomy.

Black flies (Diptera: Simuliidae) are distributed throughout the world, with more than 2200 formally described species. The family is renowned for its high frequency of cryptic species, offering an opportunity for integrative taxonomy, based on morphological, chromosomal, and molecular approaches. The biodiversity within Simulium (Psilopelmia) ignescens and S. (Psilopelmia) tunja in Colombia was estimated from the larval stage; 10 morphoforms were recognized based on 7 structural characters. This remarkable morphological variation was evaluated through 23 markers on the polytene chromosomes. We established 1 new cytoform in each nominal species. The congruence of the morphological and chromosomal assignments was evaluated using the mitochondrial marker Cytochrome Oxidase subunit I (COI) for each morphoform. The molecular data supported the chromosomal recognition of cytoforms (i.e., cryptic species). We also established the suitability of the COI marker for linking the pupal stage with each cytoform. Our results reveal the presence of hidden biodiversity in S. ignescens and S. tunja and demonstrate the power of polytene chromosomes as a tool for evaluating simuliid diversity, while illustrating the importance of integrated analyses in modern taxonomy.

Isolation of Onchocerca lupi in Dogs and Black Flies, California, USA.

In southern California, ocular infections caused by Onchocerca lupi were diagnosed in 3 dogs (1 in 2006, 2 in 2012). The infectious agent was confirmed through morphologic analysis of fixed parasites in tissues and by PCR and sequencing of amplicons derived from 2 mitochondrially encoded genes and 1 nuclear-encoded gene. A nested PCR based on the sequence of the cytochrome oxidase subunit 1 gene of the parasite was developed and used to screen Simulium black flies collected from southern California for O. lupi DNA. Six (2.8%; 95% CI 0.6%-5.0%) of 213 black flies contained O. lupi DNA. Partial mitochondrial16S rRNA gene sequences from the infected flies matched sequences derived from black fly larvae cytotaxonomically identified as Simulium tribulatum. These data implicate S. tribulatum flies as a putative vector for O. lupi in southern California.