New records for the horse fly fauna (Diptera: Tabanidae) of Jordan with remarks on ecology and zoogeography

The horse fly fauna (Diptera: Tabanidae) of Jordan is, after Israel, the richest in the Levant, with 24 known species. During the 20-year project "The Ecology and Zoogeography of the Lepidoptera of the Near East," we regularly collected blood-feeding flies, resulting in 11 additional species of Tabanidae for Jordan. The new records are: Atylotus quadrifarius (Loew, 1874), Chrysops caecutiens (Linnaeus, 1758), Dasyrhamphis nigritus (Fabricius, 1794), Haematopota pallens Loew, 1871, Nemorius irritans (Ricardo, 1901), Philipomyia graeca (Fabricius, 1794), Tabanus cordiger Meigen, 1820, Tabanus taeniola Palisot de Beauvois, 1806, Tabanus quatuornotatus Meigen, 1820, Tabanus separatus Effllatoun, 1930, and Tabanus spectabilis Loew, 1858. Most of the new records (10/11) are of Palearctic origin; of these, six are of a Mediterranean and one each of West Palearctic, Euroasiatic, Irano-Turanian, and Eremic providence. Only one species, T. taeniola, is an Afrotropical-Eremic element.     

An annotated checklist of the horse flies (Diptera: Tabanidae) of Lebanon with remarks on ecology and zoogeography: Pangoniinae and Chrysopsinae

Knowledge of the horse fly fauna (Diptera: Tabanidae) of Lebanon is fragmentary, while the local fauna of most neighboring countries has been fairly well researched. Within the framework of the 20-year project "The ecology and zoogeography of the Lepidoptera of the Near East," we regularly collected biting flies in the whole region, including Lebanon. During this time we recorded 14 horse fly species for two subfamilies in Lebanon: four Pangoniinae and ten Chrysopsinae. Only a single species, Chrysops flavipes Meigen, 1804, was known previously in Lebanon, but the following four Pangoniinae: Pangonius haustellatus (Fabricius, 1781), Pangonius obscuratus Loew, 1859, Pangonius argentatus (Szilady, 1923), and Pangonius fulvipes (Loew, 1859) and nine Chrysopsinae: Silvius appendiculatus Macquart, 1846, Silvius ochraceus Loew, 1858, Nemorius irritans (Ricardo, 1901), Nemorius vitripennis (Meigen, 1820), Chrysops buxtoniAusten, 1922, Chrysops compactusAusten, 1924, Chrysops caecutiens (Linnaeus, 1758), Chrysops italicus Meigen, 1804, and Chrysops hamatus Loew, 1858 are new records for the Lebanese fauna. The Tabanidae fauna of Lebanon is completely Palearctic and most species are of a Mediterranean distribution type. Lebanon or nearby northern Israel appears to be in the Levant, the southern geographical distribution border for the Pangoniinae and Chrysopsinae.     

Horseflies of Jordan

The horsefly (Diptera: Tabanidae) fauna of Jordan consists of 21 species belonging to seven genera. The present study adds 17 new records to this little-known group of haematophagous insects. Most of the new records were collected from several localities within the Jordan Valley. These new records are Chrysops flavipes Meigen, Atylotus farinosus (Szilády), Dasyrhamphis umbrinus (Meigen), Haematopota coronata Austen, Hybomitra decora (Loew), Hybomitra mendica (Villeneuve), Tabanus accensus Austen, T. albifacies Loew, T. bifarius Loew, T. darimonti Leclercq, T. laetetinctus Becker, T. leleani Austen, T. pallidipes Austen, T. regularis Jaennicke, T. rupinae Austen, T. sufis Jaennicke and Therioplectes tunicatus (Szilády). Tabanus albifacies and T. sufis were the most common species collected by hand nets (19.85% and 17.73%, respectively) and D. umbrinus, Haematopota minuscula Austen, and Haematopota coronata were the least common species (0.12% for each). Zoogeographical analysis of the recorded species showed that 12 species are of Mediterranean origin, four are of Afrotropical origin, two of European origin and at least three species can be considered as endemic to the Middle East.     

Anticoagulant activities in salivary glands of tabanid flies

Tabanid flies are telmophages (pool feeders), taking frequent and rapid bloodmeals from many different individual hosts. To investigate how they accomplish this intermittent feeding strategy, we examined the anticoagulant activities in salivary gland extracts (SGE) from 19 species representing six genera: Atylotus, Chrysops, Haematopota, Heptatoma, Hybomitra and Tabanus (Diptera: Tabanidae). Standard coagulation screen assays were used to determine thrombin time, prothrombin time and activated partial thromboplastin time. Chromogenic substrate assays were performed for thrombin and factor Xa activity. SGE of most species (except Chrysops spp.) considerably prolonged human plasma clotting time in a dose-dependent manner, and showed potent and specific antithrombin activity in the chromogenic substrate assay. Heptatoma pellucens displayed the strongest anticoagulant activity. Specific anti-factor Xa activity in tabanid SGE was not detected. Electrophoretic profiles of SGE proteins differed between genera and species. Overall, the results suggest that tabanids have evolved at least two antihaemostatic strategies.     

Systematics and distribution of horse flies (Diptera: Tabanidae) of Jordan.

The horse fly fauna of Jordan consists of 24 species belonging to seven genera. The present study adds two new records; Tabanus unifasciatus and Tabanus lunatus. Keys and illustrations for the horse flies of Jordan are presented based on examined materials. Distribution and geographic ranges for each species is also given.     

Vasodilatory activity in horsefly and deerfly salivary glands

Salivary gland extract (SGE) of four horsefly species (Hybomitra bimaculata Macquart, Hybomitra ciureai Séguy, Tabanus bromius L., Tabanus glaucopis Meigen) and one deerfly species (Chrysops relictus Meigen) (Diptera: Tabanidae) were shown to contain vasodilatory activity. Aliquots equivalent to 1, 5 and 10 pairs of salivary glands (SG) relaxed rat femoral artery (with intact endothelium) pre-constricted with phenylephrine. Vasodilatory activity was dose-dependent. SGE of one horsefly species (Haematopota pluvialis L.) did not induce relaxation. The kinetics of vasodilation induced by SGE of four horsefly species differed from the deerfly. These results indicate that tabanid species may produce more than one type of vasodilator to aid blood feeding.     

Distribution of halophilous species of horse flies in Croatia (Diptera: Tabanidae)

In the past 20 years, fauna and ecology of horse flies (Tabanidae) were intensively studied in Croatia, especially in the Ramsar sites as wetlands of international importance. Different habitat requirements of larvae were recognised and adults of 78 species in 10 genera were recorded. The true aquatic and semiaquatic larvae are represented by several species of the genera Chrysops Meigen, 1803 Hybomitra Enderlein, 1922. Some species from this ecological group are typical inhabitants of saline biotopes. In the Croatian fauna three halophilous species were recorded: Chrysops italicus Meigen, 1804, Hybomitra acuminata (Loew, 1858) and H. expollicata (Pandellé, 1883). The distribution of these species in Croatia covers 7, 17 and 3 UTM (Universal Transverse Mercator) grids, respectively. Specimens were primarily collected in habitats with brackish water along the Adriatic coast, such as Vransko Lake, the mouth of the River Cetina, the delta of the River Neretva and Mljet Island. Only specimens of H. acuminata were recorded in the continental part of Croatia. They were sporadically collected on localities along the Danube floodplain and on four localities near the river Drava. The distribution of H. acuminata in the continental part indicates otherwise rare saline habitats along the Drava and Danube rivers (1.38% of the total catch of the Hybomitra – species from the floodplain).     

Performance of painted plywood and cloth Nzi traps relative to Manitoba and greenhead traps for tabanids and stable flies

Experiments were conducted to adapt the cloth Nzi trap to a format suitable for fixed applications in biting fly sampling or control. Catches of tabanids [Tabanus L., Chrysops (Meigen), and Hybomitra Enderlein], and stable flies [Stomoxys calcitrans (L.)] in painted plywood traps were compared with those in standard phthalogen blue cloth traps, and in similarly painted cloth traps. The Manitoba horse fly trap and the Tabanus nigrovittatus Macquart "greenhead" box trap were used as additional standards during one tabanid season. Shiny features of traps reduced catches, e.g., paint on cloth instead of wood, or use of aluminum screening instead of netting. Nevertheless, appropriately painted plywood Nzi traps caught as many biting flies as did standard cloth Nzi traps, if paint finishes were matte, and with the use of phthalogen blue colorants. Nzi traps collected about the same tabanid fauna as the Manitoba and T. nigrovittatus traps, but with improved catches of Chrysops and Tabanus. Recommendations are provided on appropriate color matching, and selection of readily-available materials for trap construction.     

Molecular phylogeny of the horse flies: a framework for renewing tabanid taxonomy

Horse flies, family Tabanidae, are the most diverse family‐level clade of bloodsucking insects, but their phylogeny has never been thoroughly explored using molecular data. Most adult female Tabanidae feed on nectar and on the blood of various mammals. Traditional horse fly classification tends towards large heterogeneous taxa, which impede much‐needed taxonomic work. To guide renewed efforts in the systematics of horse flies and their relatives, we assembled a dataset of 110 exemplar species using nucleotide data from four genes—mitochondrial CO1, and nuclear 28S, CAD and AATS. All commonly recognized tribes in Tabanidae are represented, along with outgroups in Tabanomorpha. The phylogeny is reconstructed using Bayesian inference, and divergence times are estimated using Bayesian relaxed clock methods with time constraints from tabanid fossils. Our results show Athericidae strongly supported as the lineage most closely related to Tabanidae, and Pangoniinae and Tabaninae as monophyletic lineages. However, Chrysopsinae is nonmonophyletic, with strong support for both a nonmonophyletic Bouvieromyiini and for Rhinomyzini as sister to Tabaninae. Only the tribes Philolichini, Chrysopsini, Rhinomyzini and Haematopotini are recovered as monophyletic, although Scionini is monophyletic with exclusion of the peculiar genus Goniops Aldrich. Mycteromyia Philippi and Adersia Austen, two enigmatic genera sometimes placed in separate family‐level groups, are recovered inside Pangoniini and Chrysopsini, respectively. Several species‐rich genera are not recovered as monophyletic, including Esenbeckia Rondani, Silvius Meigen, Dasybasis Macquart and Tabanus L. Tabanidae likely originated in the Cretaceous, and all major extant groups were present by the early Palaeogene. This newly revised phylogenetic framework for Tabanidae forms the basis for a new assessment of tabanid diversification and provides context for understanding the evolution of trophic specialization in horse flies.     

Analysis of the feeding sites for some horse flies (Diptera, Tabanidae) on a human in Croatia

The landing patterns of horse flies on the human body were observed in Croatia. A total of 386 horse flies belonging to 22 species were sampled. The five most commonly collected species were used in the analysis. The stochastic linear connection is tight among the landings of the species Tabanus bromius, Tabanus maculicornis, Tabanus tergestinus, and Philipomyia graeca on the human body regions (matrix R). The preferred feeding area for these four species was the lower leg, whereas for the species Haematopota pluvialis it was the head and neck. Of the total number of horse flies that landed 44.81% were on the lower leg. Only 0.26% landed on the forearm. Chi-square analysis indicated non random landing patterns on human by these horse flies.     

Species diversity and abundance of Tabanus spp. (Diptera: Tabanidae) in different habitats of Thailand

Tabanus spp. or horse flies (Diptera: Tabanidae) are haematophagous flies of medical and veterinary importance. They are known to cause trypanosomosis or surra in domestic and wild animals in Thailand. This study conducted an entomological survey of horse flies from different sites in Thailand. Horse flies were collected from three different habitats: primary forests, secondary forests and villages using Nzi traps between April 2012 and December 2016. A total of 1835 female horse flies were collected and 45 species were identified. The five most abundant species were T. striatus (25.45%), followed by T. megalops (21.36%), T. rubidus (14.82%), T. tamthaiorum (7.90%) and T. oxybeles (6.38%). The highest proportion of horse flies was collected in villages (39.13%), followed by primary forests (34%) and secondary forests (26.87%). The species diversity of horse flies in primary forests was higher than in other habitats. The results of this study may be used for a horse fly control program.     

Seasonal abundance of horse flies (Diptera: Tabanidae) from two locations in eastern Croatia.

A total of 10,539 tabanid horse flies from 22 species and five genera was collected in the Tikves forest within the Kopacki rit Nature Park in eastern Croatia. Seasonal abundance was analyzed for the six most abundant species. Tabanus maculicornis, Tabanus tergestinus, and Haematopota pluvialis reached their highest peak abundance in the fourth week of June. Atylotus loewianus and Tabanus bromius reached their highest peak of abundance in the first week of August, whereas Tabanus sudeticus reached its maximum abundance in the third week of July. Horse flies also were collected once a week on the pasture at Petrijevci from mid-May to mid-September during 1993. Paired collections were made from a Malaise trap and from a horse by using a sweep net. A total of 2,867 tabanids belonging to 26 species was collected. The number of tabanids collected on horses was much higher than the total captured with Malaise traps. On their natural host (horse), 2.6 times more tabanids were collected than in the traps. Seasonal abundance was analyzed only for the eight most abundant species. Chrysops paralellogrammus, Tabanus autumnalis, Tabanus bromius, Tabanus tergestinus, Haematopota pluvialis, and Haematopota subcylindrica all reached their highest peak of abundance in the second week of July, whereas Tabanus maculicornis reached the maximal peak of abundance in the third week of June. Seasonal meteorological variability that occurs periodically from one year to another has a significant influence on the maximal peaks of tabanid abundance.     

Effects of horsefly (Tabanidae) salivary gland extracts on isolated perfused rat heart

The speed with which horseflies (Diptera: Tabanidae) obtain a bloodmeal suggests they have potent vasodilators. We used isolated perfused rat heart to examine the vasoactivity of salivary gland extracts (SGEs) of three horsefly species, Hybomitra bimaculata Macquart, Tabanus bromius Linnaeus and Tabanus glaucopis Meigen. Administration of horsefly SGEs to the heart produced biphasic coronary responses: a decrease and subsequent increase in coronary flow (CF), characterized by initial vasoconstriction followed by prolonged vasodilation of coronary vessels. However, although SGEs of H. bimaculata induced a significant decrease in left ventricular pressure (LVP), the effect on changes in CF was not significant except at the highest dose tested. The ability to reduce LVP without significantly lowering CF, or affecting heart rate and rhythm, represents a unique set of properties that have considerable therapeutic potential if they can be reproduced by a single molecule.     

Simplified media for spiroplasmas associated with tabanid flies

Traditionally, isolation, maintenance, and testing of Spiroplasma species (Mollicutes: Entomoplasmatales) from horse flies (Tabanus spp.) and deer flies (Chrysops spp.) (Diptera: Tabanidae) have been accomplished in the complex M1D medium. A relatively inexpensive, simplified medium for tabanid spiroplasmas could expedite procedures that require large quantities of growth medium. Nine strains of spiroplasmas, eight from tabanids and one from mosquitoes, were cultured in three simplified broth media, R2, R8-1, and C-3G, and in M1D. There was no significant difference in the rate of spiroplasma growth in M1D and the three simplified media. R2 medium supported the growth of tabanid spiroplasmas more consistently and with better morphology through 10 subcultures than did the other simplified media. Primary isolations were made in R2 medium from tabanids collected (i) in Georgia, U.S.A., with 10 isolations from 10 flies and (ii) in coastal Costa Rica, with isolation rates of 70% (28/40) and 73% (27/37), respectively, for R2 and M1D. Of the seven group VIII field isolates from Costa Rica, four were capable of sustained growth in R2, and three were triply cloned in this simplified medium. These results suggest that the simplified medium R2 is suitable for many procedures with tabanid spiroplasmas.     

Meteorological effects on the daily activity patterns of tabanid biting flies in northern Queensland,Australia

Information on the daily activity patterns of tabanid flies is important in the development of strategies that decrease the risk of pathogens transmitted by them. In addition, this information is useful to maximize numbers of tabanids trapped during short‐term studies and to target feeding behavior studies of certain tabanid species to their times of peak activity. The current study examined the effects of various meteorological factors on the daily activity patterns of common tropical species of tabanids in north Queensland. Each species studied responded differently to weather factors. Tabanus townsvilli Ricardo (Diptera: Tabanidae) was most active during late morning and early afternoon, whereas Pseudotabanus silvester (Bergroth) and Tabanus pallipennis Macquart were most active in the late afternoon. Tabanus dorsobimaculatus Macquart was most active in the morning and early afternoon. Data on daily activity patterns of tabanid flies indicates that in an area such as Townsville, North Queensland, where several species of tabanid are present concurrently in high numbers, the overlapping periods of high activity for these species indicate a high risk of pathogen transmission for most of the day (10.00–19.00 hours). Similarly, because each species responds differently to weather variables, only extreme weather conditions are likely to inhibit activity of all species. These data also indicate that for maximal results, trapping and feeding behavior studies should be tailored to the preferred activity period of the species under investigation.     

ACTION DE L'ANHYDRIDE CARBONIQUE ET DE FACTEURS VISUELS SUR LES PERFORMANCES DE PIEGES «MANITOBA» MODIFIES EN MILIEU MONTAGNARD

Résumé Les performances de pièges à Tabanidae de type Manitoba ont été étudiées en milieu montagnard et subalpin. On a testé les influences respectives de la présence ou de l'absence d'une source de CO₂, de la nature de cette source (solide ou gazeuse), de la présence ou de l'absence d'un leurre visuel (sphère noir brillant), de la nature transparente ou opaque de la partie supérieure du piège (hotte conique). Plus de 8.000 Tabanidae et plus de 51.000 Muscidae ont été pris au cours de ces essais totalisant 660 heures-pièges.

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Summary Manitoba traps, traps without decoy or with black canopy, unbaited or baited with dry ice or compressed carbon dioxide, were compared for the numbers of horse flies and muscid flies caught in mountainous mixed wood and pasture land and in subalpine larch forest between 1500 m and 2200 m, in July. More muscid flies were captured in the compressed CO₂ baited traps than in dry ice baited traps. Haematopota pluvialis, Tabanus maculicornis and Tabanus quatuornotatus were more attracted by CO₂ than Tabanus bromius. T. bromius was more attracted by the decoy than other species. The genus Atylotus and T. bromius are more attracted to the black canopy than other species. The visual decoy is more necessary to the efficiency of the trap than the chemical bait. The CO₂ baited Manitoba trap caught the greatest number of horse flies (180 per trap and per hour). The unbaited Manitoba trap was efficient too, but other traps were inefficient. About 8000 horse flies and 51000 muscid flies were caught.

     

Bloodsucking dipteran insects (Diptera) attacking humans and animals (the “gnus” complex) in northwestern Russia: I. General characteristics of the fauna

The fauna of bloodsucking Diptera of northwestern Russia comprises 226 species of mosquitoes (Culicidae), biting midges (Ceratopogonidae), black flies (Simuliidae) and horseflies (Tabanidae). In particular, Culicidae include 41 species from 10 subgenera of 5 genera of 2 subfamilies. Simuliidae are represented by 110 species of 23 genera, belonging to 6 tribes of 2 subfamilies; the genus Culicoides of the Ceratopogonidae, by 31 species; and Tabanidae, by 44 species of 7 genera from 2 subfamilies. There is a necessity of elaboration and application of more advanced forms of information storage and field data standardization. In the Zoological Institute, Russian Academy of Sciences, a special database on the bloodsucking insects of Russia is created in the DBMS Access. A list of abundant bloodsucking species of mosquitoes, biting midges, black flies and horseflies is included. A brief synopsis of bibliography of investigations of the bloodsucking dipterans from northwestern Russia is given, provided with a list of more then 190 publication.     

Seasonal surveillance of deer and horse flies (Diptera: Tabanidae), Gyeonggi province,Republic of Korea

Deer and horse flies, Family Tabanidae, are biological, mechanical, and potential vectors of approximately 27 viral, bacterial, and helminthic pathogens of veterinary and medical importance. Biting fly surveillance was conducted using New Jersey light traps and Mosquito Magnet® traps from May-October during 2015–2018 at eight sites (villages and US military installations and training areas) near/in the Demilitarized Zone and at a US military installation in southern Gyeonggi province, Republic of Korea.A total of 9985 deer and horse flies, comprising 4 genera and 17 species, were collected. The predominant species collected was Chrysops mlokosiewiczi (95.39%), followed by Chrysops suavis (1.48%) and Haematopota koryoensis (1.07%), while the remaining species accounted for only 2.06% of the specimens collected. A bimodal peak was observed for C. mlokosiewiczi in June and late-July, but not for the other two predominant species during the whole of the study period. This study provides and insight into ecological behavior and seasonal abundance of deer and horse flies in Gyeonggi province, Republic of Korea.     

Feeding strategies of the horseflies Hybomitra expollicata and Tabanus bromius in southern France

Blood-feeding success of female horseflies, Hybomitra expollicata Pandellé and Tabanus bromius L. (Diptera: Tabanidae), was studied. Bloodmeal weights of flies permitted different periods of residence on a horse's belly were estimated, and related to subsequent ovarian development and the number of mature eggs produced. Fitness gain per visit to a host (as estimated by the number of mature eggs produced) increased more rapidly with residence time on the host for H.expollicata than for T.bromius. H.expollicata inhibed blood more rapidly and, above a threshold bloodmeal size, produced a consistently higher number of eggs per unit weight of blood ingested. The feeding strategies of the two species are compared.     

The Horsefly Fauna (Diptera,Tabanidae) of Yakutia

The literature on the fauna of the family Tabanidae (Diptera) of Yakutia is analyzed. The distribution of 37 species of horseflies over 7 natural regions of Yakutia is considered. Seven taxa of horseflies previously known by erroneous records are excluded from the fauna of Yakutia.