Macaronesian Muscidae (Diptera). IV. The genus Helina Robineau-Desvoidy, and description of the male of Spilogona maderensis Michelsen.

The Macaronesian species of the muscid genus Helina are revised. The genus is represented in the Canary Islands by 9 species, in Madeira by 7 species, and in the Azores Islands by 1 species. Four species, one divided in two subspecies, are endemic to the Canary Islands: Helina grancanariae sp. nov. (Gran Canaria), H. obscurisquama obscurisquama (Stein) (Tenerife, La Gomera), H. obscurisquama pseudonivalis (Santos Abréu) ssp. rev. (La Palma), H. rufitibialis Hennig (Tenerife, La Gomera, La Palma) and H. santosabreui sp. nov. (La Gomera, La Palma). The following three species are endemic to Madeira: Helina atlantica (Tiensuu), H. lundbladi Tiensuu and H. vilissima Hennig. Existing records of Helina almeriensis (Strobl) and H. quadrum (Fabricius) from the Canary Islands are considered as misidentifications. An identification key to males and females concludes the species accounts. Finally, a description is given of the previously unknown male sex of the endemic Madeiran Spilogona maderensis Michelsen.

An updated key to the genus Limnophora Robineau-Desvoidy (Diptera: Muscidae) from China, with descriptions of two new species.

A key to the 80 species and 2 subspecies of the genus Limnophora Robineau-Desvoidy, 1830 from China based on males is provided, including two new species Limnophora magnipulvilla sp. nov. and Limnophora apiciflata sp. nov. Photographs of habitus and male genitalia as well as illustrations of the male genitalia of the new species are given.

How strange: Coenosia curiosa sp. nov. (Diptera: Muscidae), the first recorded Tiger fly from Lesotho, with revision of the Coenosia globuliseta-group.

A new species of tiger fly, Coenosia curiosa sp. nov., is described from the Kingdom of Lesotho. It is also the first record of the genus for Lesotho. An identification key is provided for the C. globuliseta-group, which now includes four species with aberrant males that have enlarged apically globular frontal setae, thus far known only from southern Africa. Diagnoses, material examined and terminalia illustrations for all known species of the C. globuliseta-group are also provided.

Avian vampire fly (Philornis downsi) mortality differs across Darwin's finch host species.

In invasive parasites, generalism is considered advantageous during the initial phase of introduction. Thereafter, fitness costs to parasites, such as host-specific mortality, can drive parasites towards specialism to avoid costly hosts. It is important to determine changes in host specificity of invasive populations to understand host-parasite dynamics and their effects on vulnerable host populations. We examined changes in mortality in the introduced avian vampire fly (Philornis downsi) (Diptera: Muscidae), a generalist myasis-causing ectoparasite, between 2004 and 2020 on Floreana Island (Galápagos). Mortality was measured as the proportion of immature larvae found upon host nest termination. Over the time period, the avian vampire fly was most abundant and had low mortality in nests of the critically endangered medium tree finch (Camarhynchus pauper) and had the highest mortality in nests of hybrid tree finches (Camarhynchus spp.). Low larval mortality was also found in small tree (Camarhynchus parvulus) and small ground finch (Geospiza fuliginosa) nests. Selection could favour avian vampire flies that select medium tree finch nests and/or avoid hybrid nests. Overall, the finding of differences in avian vampire fly survival across host species is parsimonious with the idea that the introduced fly may be evolving towards host specialisation.

Wing morphometrics of medically and forensically important muscid flies (Diptera: Muscidae).

Many muscid flies (Diptera: Muscidae) are well-known as medical, veterinary, and forensically significant insects, thus correct species identification is critically important before applying for fly control and determining a minimal postmortem interval (PMImin) in forensic investigations. Limited in taxonomic keys and taxonomists, as well as scanty in advanced molecular laboratories lead to difficulty in identification of muscids. To date, a landmark-based geometric morphometric analysis of wings has proven to be a promising alternative technique for identifying many insect species. Herein, we assessed wing morphometric analysis for identification of six medically and forensically important muscids, namely Musca domestica Linnaeus, Musca pattoni Austen, Musca ventrosa Wiedemann, Hydrotaea chalcogaster (Wiedemann), Hydrotaea spinigera Stein, and Dichaetomyia quadrata (Wiedemann). A total of 302 right wing images were digitized based on 15 homologous landmarks and wing shape variation among genera and species was analyzed using canonical variate analysis, whereas sexual shape dimorphism of M. domestica, M. ventrosa, and D. quadrata was analyzed using discriminant function analysis. The cross-validation revealed a relatively high percentage of correct classification in most species, ranging from 86.4% to 100%, except for M. pattoni, being 67.5%. Misidentifications were mainly due to cross-pairings of the genus Musca; M. domestica VS M. pattoni VS M. ventrosa. The accuracy of classification using cross-validation test demonstrated that wing shape can be used to evaluate muscid flies at the genus- and species-level, and separate sexes of the three species analyzed, with a high reliability. This study sheds light on genus, species, and sex discrimination of six muscid species that have been approached using wing morphometric analysis.

Macaronesian Muscidae (Diptera). II. The genus emLimnophora/em Robineau-Desvoidy with description of a new Canarian endemic species.

The Macaronesian species of the muscid genus Limnophora are reviewed with special emphasis on the fauna of the western Canary Islands. The genus is represented by 14 species in the Macaronesian archipelagos, with 10 species in the Canary Islands, 4 species in the Cape Verde Islands, and 3 species in Madeira. Limnophora obsignatula sp. nov. is endemic to the western Canary Islands, where it replaces the widespread continental L. obsignata (Rondani). It is further shown that L. paneliusi Emden, a species so far considered endemic to the Cape Verde Islands, is widespread in the Canary Islands. Two species, L. obsignata (Rondani) and L. tigrina Am Stein, are removed from the list of Canarian Muscidae as based on misidentifications of other species with a mesonotal "Anthomyia-pattern". Limnophora (Calliophrys) riparia capoverdica Emden, a taxon described from the Cape Verde Islands, is synonymized with L. riparia (Fallén), syn. nov. Distribution data and illustrated diagnoses are given for each species. The species account is finalized with an identification key to males and females. An assessment of the ovipositor as a characters source in phylogeny and species recognition is made for 12 species of Limnophora. Finally, it is pointed out that several species are acutely threatened due to habitat disturbance.

Macaronesian Muscidae (Diptera). III. First record of the genus Spilogona Schnabl based on two new endemic species from the Azores and Madeira.

The occurrence of the muscid genus Spilogona Schnabl in Macaronesia is documented for the first time by the description of two new species, S. azorensis sp. nov. and S. maderensis sp. nov. These appear to be local endemics of the Azores and Madeira, respectively.

Review of Synthesiomyia nudiseta (Diptera: Muscidae) as a useful tool in forensic entomology.

Synthesiomyia nudiseta (van der Wulp, 1883) is a synanthropic muscid found in tropical and subtropical zones around the world. The larvae of this species are a secondary agent of myiasis with necrophagous habits and play an important role in forensic entomology, as they are used as an indicator of post-mortem interval. Adults can be considered vectors of etiological agents such as Escherichia coli and Shigella dysenteriae. Due to its ability to adapt to different environmental conditions, its high dispersal capacity (shown by its introduction to Europe), its predatory habits in the last larval stage and the difficulty of identifying it, a very important goal is to update our knowledge about this species. Therefore, the main objective of this paper is to review the identification, geographical distribution and biology of this species in order to provide better support to investigations involving this fly.

More than just nestlings: incidence of subcutaneous Philornis (Diptera: Muscidae) nest flies in adult birds.

Philornis flies Meinert (Diptera: Muscidae) have been documented parasitizing over 250 bird species, some of which are endemic species threatened with extinction. Philornis parasitism is hypothesized to affect nestlings disproportionately more than adult birds because limited mobility and exposed skin of nestlings increase their vulnerability to parasitism. We used a comprehensive literature review and our recent fieldwork in the Dominican Republic, Puerto Rico, and Grenada to challenge the idea that parasitism by subcutaneous Philornis species is a phenomenon primarily found in nestlings, a fact that has not been quantified to date. Of the 265 reviewed publications, 125 (49%) reported incidences of parasitism by subcutaneous Philornis, but only 12 included the sampling of adult breeding birds. Nine of these publications (75%) reported Philornis parasitism in adults of ten bird species. During fieldwork in the Dominican Republic, Puerto Rico, and Grenada, we documented 14 instances of parasitism of adult birds of seven avian species. From literature review and fieldwork, adults of at least fifteen bird species across 12 families and four orders of birds were parasitized by at least five Philornis species. In both the published literature and fieldwork, incidences of parasitism of adult birds occurred predominantly in females and was frequently associated with incubation. Although our findings indicate that Philornis parasitism of adult birds is more common than widely presumed, parasite prevalence is still greater in nestlings. In the future, we recommend surveys of adult birds to better understand host-Philornis relationships across life stages. This information may be essential for the development of effective control measures of Philornis to ensure the long-term protection of bird species of conservation concern.

Ethiopian muscids (Diptera, Muscidae) egg-carriers of Stylogaster Macquart (Diptera, Conopidae).

We here report for the first time on the presence of three species of the conopid genus Stylogaster Macquart (Diptera, Conopidae) in Ethiopia, viz. S. nitens Brunetti, S. westwoodi Smith and Stylogaster sp.. We further screened 908 muscid flies (Diptera, Muscidae) for the presence of impaled eggs of Stylogaster and recorded eggs on 89 individuals (9.8%). Eggs were impaled on eight species, viz. Limnophora translucida Stein, Musca lusoria Wiedemann, Musca splendens Pont, Neomyia chrysopyga (Emden), Pseudohelina nigritarsis (Jaennicke), Stomoxys omega Newstead, Stomoxys taeniatus Bigot and Stomoxys varipes (Bezzi). The maximum number of eggs found on a single muscid was six. We illustrated the dissected eggs. L. translucida, M. lusoria, M. splendens, N. chrysopyga and S. varipes are reported as new muscid hosts species for Stylogaster.

First record of Fanniidae (Diptera) from New Caledonia.

The family Fanniidae is newly recorded to New Caledonia. Adult male and female of Fannia capitalis Pont are illustrated as well as the peculiar male terminalia, the female ovipositor, and the egg. Male phallic process and egg are described for the first time.

Characterization and Identification of Puparia of Hydrotaea Robineau-Desvoidy, 1830 (Diptera: Muscidae) From Forensic and Archaeological Contexts.

Flies and beetles are the main components of the entomofauna colonizing a body after death. Following the recognition of constant and predictable colonization patterns and the knowledge about the dependence of the insect development to temperature, a new discipline, forensic entomology, has provided information useful to reconstruct criminal events. Funerary archaeoentomology has also applied the same rationale in archaeological contexts. Puparia represent a large fraction of the insect remains that can be found associated with a cadaver, especially when the body is mummified or in the advanced stages of decomposition. Puparium identification is still a problematic topic due to the lack of identification keys and, in several cases, a lack of diagnostic feature descriptions. Here, we focus the attention on some Hydrotaea Robineau-Desvoidy (Diptera, Muscidae) puparia from forensic and archaeological contexts. Puparia of Hydrotaea capensis (Wiedemann), Hydrotaea ignava (Harris), Hydrotaea aenescens (Wiedemann), Hydrotaea similis Meade, Hydrotaea pilipes Stein, and Hydrotaea dentipes (Fabricius) are here detailed and illustrated. Posterior spiracles, anal plate, and intersegmental spines have been considered as good diagnostic characters for the identification of these puparia.

A review and illustrated description of Musca crassirostris, one of the most neglected haematophagous livestock flies.

Tabanids, stomoxyine flies, hippoboscids and tsetse flies are the most well-known brachyceran biting flies of livestock. Only a few other higher Diptera have developed the unique mouthparts required for blood feeding. These neglected blood feeders can also have direct effects on hosts through blood loss, and are likely to contribute to the transmission of pathogens. Musca crassirostris (Diptera: Muscidae) is one of the most abundant of the muscid flies with this haematophagous lifestyle; it is widespread in the Palaearctic, Afrotropical and Oriental regions. The present study reviews and summarizes the biology and morphology of this species, and its potential for impact on animals and humans. The study also provides a fully illustrated description of the fly to facilitate its identification, and reviews information on abundance, with a focus on recent trapping surveys in Thailand. When sampled using traps designed for other biting flies, M. crassirostris appears to be four and 45 times more abundant than stomoxyines and tabanids, respectively. High numbers of M. crassirostris in the vicinity of livestock have also been associated with outbreaks of disease, such as that of a fatal plague in bovine farms in Egypt. This calls for a reconsideration of its potential impacts on livestock economics and health, and thus the development of suitable control methods.

Phenotypic susceptibility to pyrethroids and organophosphate of wild Stomoxys calcitrans (Diptera: Muscidae) populations in southwestern France.

Stomoxys calcitrans (Diptera: Muscidae), is an important vector of lumpy skin disease and bovine besnoitiosis in Europe. Control of this biting fly could represent a keystone in the containment of this emerging disease. Reports of insecticide resistance in S. calcitrans are scarce in Europe. Therefore, the purpose of this study was to evaluate the phenotypic susceptibility to deltamethrin, cypermethrin and phoxim of five wild S. calcitrans populations from southwestern France, where transmission of bovine besnoitiosis is very prevalent. Adult S. calcitrans were caught at each study site and exposed to insecticide-impregnated filter papers under laboratory conditions. Quantities of active ingredients on filter papers corresponded to the recommended doses proposed by the manufacturers (37.5 mg a.i./m2 of cattle's skin, 125 mg a.i./m2 and 750 mg a.i./m2 for deltamethrin, cypermethrin and phoxim respectively) were tested. Knock-down effects (KD) (1 h after the onset of exposure) and mortality rates (24 h and 48 h after exposure) were evaluated. Phoxim showed a rapid and full efficacy in all populations. However, the KD effects (37.5 to 97.5%) and the mortality rates at 48 h (10 to 91.25%) induced by the exposure to pyrethroids varied greatly according to the study site but none of the populations showed full susceptibility. Therefore, the current recommended doses of these pyrethroids are probably less efficient than expected in the field and should be considered with caution in the control of bovine besnoitiosis in France.

The Application of COI Gene for Species Identification of Forensically Important Muscid Flies (Diptera: Muscidae).

Muscid Flies (Diptera: Muscidae) are of great forensic importance due to their wide distribution, ubiquitous and synanthropic nature. They are frequently neglected as they tend to arrive at the corpses later than the flesh flies and blow flies. Moreover, the lack of species-level identification also hinders investigation of medicolegal purposes. To overcome the difficulty of morphological identification, molecular method has gained relevance. Cytochrome c oxidase subunit I (COI) gene has been widely utilized. Nonetheless, to achieve correct identification of an unknown sample, it is important to survey certain muscid taxa from its geographic distribution range. Accordingly, the aim of this study is to contribute more geographically specific. We sequenced the COI gene of 51 muscid specimens of 12 species, and added all correct sequences available in GenBank to yield a total data set of 125 COI sequences from 33 muscid species to evaluate the COI gene as a molecular diagnostic tool. The interspecific distances were extremely high (4.7-19.8%) in either the standard barcoding fragment (658 bp) or the long COI sequence (1,019-1,535 bp), demonstrating that these two genetic markers were nearly identical in the species identification. However, the intraspecific distances of the long COI sequences were significantly higher than the barcoding region for the conspecific species that geographical locations vary greatly. Therefore, genetic diversity presented in this study provides a reference for species identification of muscid flies. Nevertheless, further investigation and data from more muscid species are required to enhance the efficacy of species-level identification using COI gene as a genetic marker.

Insects Associated with Pig Carrion in Two Environments of the Brazilian Savanna.

Studies of carrion fauna have increased in Brazil and have contributed to the knowledge of this fauna in the national territory. Brazil has continental dimensions and presents various biomes in its territorial area. Most of the carrion fauna have seldom been studied, and this is especially true for the Savanna or "Cerrado." The present research examined the fauna visiting carrion in two environments and two seasons in a rural area situated in a region of Cerrado in southeastern Brazil. The two environments studied were a pasture and a fragment of semi-deciduous forest. Samples were collected during the dry and humid seasons of the year. The study identified the diversity and relative abundance of species attracted to pig carcasses (Sus scrofa L.) exposed in each environment and season. Eight pig carcasses (10 ± 1 kg) were placed in traps, and adult insects attracted to the traps were collected during the decay of the carcasses. A total of 92,489 insects were collected during the experiments. The Diptera was the most frequent order (92.2%) represented by 27 families and at least 124 species. This was followed by Coleoptera (4.4%) represented by 14 families and at least 65 species. Sarcophagidae had the greatest diversity of species, followed by Muscidae and Calliphoridae. Several species were collected only during a specific period of the year or in a single environment. The results are important from an ecological point of view and for Medico-Legal Forensic Entomology.

Identification of Muscidae (Diptera) of medico-legal importance by means of wing measurements.

Cadavers attract numerous species and genera of Muscidae, both regular elements of carrion insect assemblages, and accidental visitors. Identification of adult Muscidae may be considered difficult, particularly by non-experts. Since species identification is a vital first step in the analysis of entomological material in any forensic entomology orientated experiment and real cases, various alternative methods of species identification have been proposed. We investigated possibility of semiautomated identification by means of wing measurements as an alternative for classic morphology and DNA-based approaches. We examined genus-level identification success for 790 specimens representing 13 genera of the most common European cadavers visiting Muscidae. We found 99.8% of examined specimens correctly identified to the genus-level. Without error, the following were identified: Azelia, Eudasyphora, Graphomya, Hydrotaea, Musca, Muscina, Mydaea, Neomyia, Polietes, Stomoxys and Thricops. Genus-level misidentifications were found only in Helina and Phaonia. Discrimination of examined material on the species level within Hydrotaea (318 specimens representing eight species) and Muscina (163 specimens representing four species) showed lower, yet still high average identification success, 97.2 and 98.8%, respectively. Our results revealed relatively high success in both genus and species identification of Muscidae of medico-legal importance. Semiautomated identification by means of wing measurements can be used by non-experts and does not require sophisticated equipment. This method will facilitate the identification of forensically relevant muscids in comparison to more difficult and more time-consuming identification approaches based on taxonomic keys or DNA-based methods. However, for unambiguous identification of some taxa, we recommend complementary use of identification keys.

Afrotropical Limnophora Robineau-Desvoidy (Diptera, Muscidae) with the description of four new species.

All the Afrotopical Limnophora Robineau-Desvoidy (Diptera, Muscidae) deposited in the Natural History Museum, London (United Kingdom) were analyzed, including the types of the species described by Emden, mainly from the British Museum (Natural History) Ruwenzori Expedition of 1934-1935. Diagnoses and notes on all the species recorded from the Afrotropical region are given, together with illustrations of the male terminalia (when sufficient material was available), some of which are given for the first time. We made the following taxonomic changes: Limnophora terrestris Paterson, 1955 is a junior synonym of Limnophora translucida Stein, 1913 (syn. nov.); Limnophora aculeipes latilamellata Malloch, 1921 has its status revised and is elevated to species rank (stat. rev.) with Limnophora aculeipes eurymetopa Emden, 1951 proposed as a junior synonym (syn. nov.); L. majuscula kinangopana Emden, 1951 is also raised to species rank (stat. nov.). Four new species are described: Limnophora alta sp. nov.; Limnophora nuda sp. nov.; Limnophora numerosa sp. nov. and Limnophora sinuosa sp. nov., with illustrations of male terminalia.

Landmark and outline-based geometric morphometrics analysis of three Stomoxys flies (Diptera: Muscidae).

Adult flies of the genus Stomoxys Geoffroy, 1762 (Diptera: Muscidae), especially S. pullus Austen, 1909, S. uruma Shinonaga et Kano, 1966 and S. indicus Picard, 1908, are morphologically similar and sometimes difficult to distinguish when using external morphological characteristics. These species may act as vectors and/or potential vectors of many pathogens (virus, bacteria and protozoa). Their correct identification is important to target the vectors involved in the transmission of the pathogens and also helps in the fly control program.The aim of the present study was to distinguish three species which are difficult to separate using traditional diagnostic characters for species of Stomoxys such as colour patterns and body proportions. Modern morphometrics, both landmark and outline-based, was used to access wing geometry of S. pullus, S. uruma and S. indicus. A total of 198 and 190 wing pictures were analysed for landmark- and outline-based approaches, respectively. Wing shape was able to separate species and sexes of the three Stomoxys flies with highly significant difference of Mahalanobis distances. The cross-validated classification scores ranged from 76% to 100% for landmark and 77% to 96% for outline-based morphometrics. The geometry of wing features appears to be a very useful, low-cost tool to distinguish among the vectors S. pullus, S. uruma and S. indicus.

Taxonomy of Morellia Robineau-Desvoidy (Diptera: Muscidae): revision of the subgenera Morellia s. str. and Parapyrellia Townsend.

The subgenera Morellia Robineau-Desvoidy, 1830 and Parapyrellia Townsend, 1915 of Morellia (Diptera, Muscidae) are revised. Forty-two species of the subgenus Morellia (out of 50) and seven of the subgenus Parapyrellia (all species) are redescribed and illustrated, and 48 species are keyed (41 spp. of Morellia and 7 spp. of Parapyrellia). Morellia (M.) cashmirana van Emden, 1965, originally described as a subspecies of M. (M.) hortorum (Fallén, 1817), is raised to species rank, stat. nov. Four species are transferred from the subgenus Morellia to the subgenus Parapyrellia (subgen. comb. nov.): M. dendropanacis Pamplona & Couri, 1995; M. lopesae Pamplona, 1986; M. roppai Pamplona, 1986 and M. semimarginata (Stein, 1918).