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Tan, D. S. H., Ang, Y., Lim, G. S., Ismail, M. R. B. & Meier, R. (2010). From ‘cryptic species’ to integrative taxonomy: an iterative process involving DNA sequences, morphology, and behaviour leads to the resurrection of Sepsis pyrrhosoma (Sepsidae: Diptera). —Zoologica Scripta, 39, 51–61. The increased availability of DNA sequences has led to a surge of ‘cryptic species’ in the literature. These units are usually proposed based on finding genetically distinct lineages within species that were initially defined based on morphological characters. However, few authors attempt to confirm whether these ‘cryptic’ units are species and even fewer authors are explicit about which species concept is applied. Here, we use an example from Sepsidae (Diptera) to demonstrate how cryptic species can be validated by an iterative process involving several data sources and an evaluation of the data under different species concepts. A phylogeographic analysis based on 50 specimens for five species of the flavimana group revealed deep mitochondrial splits within Sepsis flavimana which was suggestive of a cryptic species. We resolve the initial conflict between DNA sequences and morphology by adding new morphological data as well as behavioural evidence and tests for reproductive isolation. One cryptic species is confirmed and Sepsis pyrrhosoma, a former synonym of S. flavimana, is here shown to be a valid species under most species concepts. We can thus document that the same data can lead to similar conclusions under conflicting concepts once different kinds of data are integrated.

Fluctuating asymmetry (FA) is a widely employed measure of developmental stability. It has been found to increase with many stressors including parasite infection. Associations between parasites and FA may exist for several reasons in addition to parasites being the direct cause of increased FA. Developmentally stable individuals may have superior immune systems, and be less susceptible to parasite infection, and/or may be less exposed to parasites than developmentally unstable ones. Mites negatively impact host fitness in a number of insects, and if FA is a reflection of general genetic quality, as has been proposed, associations between mite number and FA are predicted. Potential relationships were investigated between an ectoparasitic mite, Pediculoides mesembrinae (Canestrini) (Phthiraptera: Menoponidae) and FA in the common dung fly Sepsis cynipsea (L.) (Diptera: Sepsidae). While it was found that mite infested flies died much faster than flies without mites, indicating that mites indeed stress their hosts, counter to expectations, no associations between mites and FA were found in any analyses. Additionally, FA in mite-infected flies generally did not differ from previously published FA data from uninfected S. cynipsea. Nevertheless, parasitized males tended to be somewhat less asymmetrical than non-parasitized males, but based on our data, it does not appear that mite infestation is generally associated with developmental stability in S. cynipsea.

I herein revise the genus Toxopoda Macquart of the Oriental and Australasian regions. A total of 16 species is discussed. Six new species are described: T. cavata sp. nov., T. angulata sp. nov., T. elephantina sp. nov., T. zuskai sp. nov., T. malayana sp. nov., and T. ozerovi sp. nov. The species T. contracta (Walker), T. viduata (Thomson) and T. simplex Iwasa, which can be easily confused with other species, are correctly redescribed and illustrated. Distributional notes and a key to the species of the Oriental and Australasian regions are provided.

Made available in DSpace on 2013-08-12T18:23:01Z (GMT). No. of bitstreams: 0 Previous issue date: 2008-01-01 Made available in DSpace on 2013-09-30T18:40:11Z (GMT). No. of bitstreams: 0 Previous issue date: 2008-01-01 Submitted by Vitor Silverio Rodrigues (vitorsrodrigues@reitoria.unesp.br) on 2014-05-20T13:53:05Z No. of bitstreams: 0 Made available in DSpace on 2014-05-20T13:53:05Z (GMT). No. of bitstreams: 0 Previous issue date: 2008-01-01 The decomposition of small carcasses in the open is frequently neglected although it may provide information of forensic importance. This paper describes an experimental study of arthropod species associated with carcasses of mouse, Mus musculus (Linnaeus, 1758) and rat, Rattus norvegicus (Berkenhout, 1769) (Rodentia: Muridae). Four carcasses were left inside iron cages in sunlit and shady areas in a secondary forest in Southeastern Brazil twice a season for four seasons (n = 16 carcasses of each rodent). The carcasses were removed when arthropods ceased to visit them. The visiting and colonizing invertebrates were collected daily and identified. Immatures were also collected and reared in a laboratory for identification. We collected 6,514 arthropods (820 adults and 5,694 juvenile forms) belonging to 53 species from the families Sarcophagidae, Calliphoridae, Muscidae, Fanniidae, Syrphidae, Richardiidae, Sepsidae, Micropezidae, Otitidae, Drosophilidae, Phoridae, Dolichopodidae, Anthomyiidae, Asilidae and Lauxaniidae (Diptera), Formicidae, Ichneumonidae, Encyrtidae and Apidae (Hymenoptera), Staphylinidae (Coleoptera) and Gonyleptidae (Opiliones). Lucilia eximia (Wiedemann, 1819) (Diptera: Calliphoridae) and Peckia (Pattonella) intermutans (Walker, 1861) (Diptera: Sarcophagidae) deserve special attention because both adult and immature forms were collected in all seasons and in both areas. Our results indicate that the frequency of occurrence of these arthropods was positively associated with carcass size (mouse or rat); no marked insect succession on the carcasses occurred; and the diversity of Calliphoridae and Sarcophagidae was high, irrespective of season. Univ Estadual Campinas, Inst Biol, Dept Parasitol, Campinas, SP, Brazil Univ Estadual Paulista, Inst Biociencias, Dept Parasitol, Botucatu, SP, Brazil Univ Estadual Paulista, Inst Biociencias, Dept Biol, Rio Claro, SP, Brazil Univ Estadual Paulista, Inst Biociencias, Dept Parasitol, Botucatu, SP, Brazil Univ Estadual Paulista, Inst Biociencias, Dept Biol, Rio Claro, SP, Brazil

The males of almost all sepsid species have strongly modified forelegs that are used to clamp the female's wingbase during mounting. Here, we describe a new species in the genus Perochaeta whose males have unmodified forelegs. We use DNA sequence data for ten genes to reconstruct the position of Perochaeta on the phylogenetic tree for Sepsidae, and reveal that the lack of foreleg armature in Perochaeta dikowi sp.n. is secondary. Through the study of the mating behaviour of the new species, we demonstrate that the loss of armature is correlated with a new mounting technique during which the males of P. dikowi do not use the foreleg to clamp the female's wingbase. Instead, the male approaches the female from behind and bends his abdomen forwards in order to establish genital contact. Our study shows how data from morphology, phylogenetics, and behavioural biology can complement each other to yield a deeper understanding of how changes in morphology and behaviour are correlated.

The flies of the genus Dicranosepsis from Vietnam were investigated and classified taxonomically. Six new species (D. longa sp. nov., D. kurahashii sp. nov., D. monoseta sp. nov., D. sinuosa sp. nov., D. barbata sp. nov., and D. vietnamensis sp. nov.) are described and illustrated. Dicranosepsis is redefined and a revised key to the species is also provided.

The complex, species-specific foreleg armature in males of the genus Themira (Diptera: Sepsidae) provides an ideal system for testing competing hypotheses for the evolution of sexually dimorphic character divergence. In sepsid flies, the male holds onto the female by clasping her wing base with his modified forelegs. In the present study, we document the male leg and the female wing morphology using scanning electron microscopy and confocal microscopy. We use a phylogenetic tree for Themira to reconstruct male foreleg and female wing evolution and demonstrate that the male legs have evolved elaborate structures with little or no corresponding changes in wing morphology. This lack of interspecific variation in female wings is not in agreement with the hypothesis of a morphological ‘evolutionary arms race’ between males and females. However, there is also no evidence for sex-specific wing differences in sensory organs on the wing base that may explain how females could assess males according to Eberhard's ‘cryptic female choice’ hypothesis. Finally, our study reveals the function of several novel morphological clasping structures and documents that the male foreleg characters in Themira are highly homoplastic. Male forelegs in two clades evolve considerably faster than in other species or clades. These two clades include Themira superba and Themira leachi, species that have some of the most dramatically modified forelegs known in Diptera. © 2008 The Linnean Society of London, Biological Journal of the Linnean Society, 2008, 93, 227–238.

SUMMARY Abdominal appendages in male sepsid flies are a complex novel structure of unknown developmental and evolutionary origin. Although these abdominal appendages superficially resemble serially homologous insect appendages, they do not develop from imaginal discs like other dipteran appendages. Cauterization of the genital disc and ventral abdominal histoblasts in Themira biloba (Sepsidae, Diptera) revealed that these abdominal appendages develop from the ventral histoblast nests of the fourth abdominal segment. Cell counts of the histoblasts in males and females revealed that the ventral histoblast nests on the fourth abdominal segment in males were significantly larger than other histoblast nests, indicating that the specification of that segment as the location of the abdominal appendages occurs before the last larval instar. The recruitment of histoblasts to produce appendages has not been documented before, and implies a developmental and evolutionary potential for histoblasts that was previously unknown.

The stages of decomposition and the faunal succession on rabbit carcasses in three different habitats, namely jungle, rural, and highland areas, were studied. Three New Zealand White rabbit (Oryctolagus cuniculus) carcasses weighing ∼2 kg were sampled daily until the decomposition process was completed. Representative specimens of adult flies, larvae, pupa, and mites were collected from the carcasses and processed in the laboratory. There were differences in decomposition rate and faunal succession between the carcasses. The fastest rate of decomposition was recorded in rural area, and the slowest rate of decomposition was recorded in highland area. The carcasses exhibited the same pattern of colonization by adult flies, but the dominant species of larvae and adult flies on each carcass in specific habitats were different. The primary species of flies recorded in jungle were Chrysomya megacephala F., Chrysomya rufifacies (Macquart), Chrysomya chani Kurahashi, Chrysomya villenuevi Patton, Chrysomya nigripes Aubertin, Chrysomya pinguis (Walker), Hemipyrellia ligurriens (Wiedemann), Hemipyrellia tagaliana (Bigot), Hypopyiopsis fumipennis (Walker), Hypopygiopsis violacea (Macquart), and Hydrotaea spinigera Stein represented by both adults and larvae. Musca domestica L., Atherigona sp., Lioproctia pattoni (Senior-White), Lioproctia saprianovae Pape & Banziger, and Seniorwhitea princeps (Wiedemann) were represented by adults only. The biodiversity of flies in the rural area were C. megacephala, C. rufifacies, H. ligurriens, Fannia canicularis L., Hydrotaea chalcogaster (Wiedemann), and Hyd. spinigera represented by both adults and larvae, meanwhile M. domestica, Atherigona sp., Boettcherisca peregrina (Robineau-Desvoidy), Parasarcophaga taenionota Wiedemann, Parasarcophaga scopariiformis Senior-White, and S. princeps were represented by adults only. The species of flies collected in the highland area were Lucilia porphyrina (Walker), C. megacephala, C. rufifacies, C. villenuevi, C. pinguis, H. ligurriens, Hyd. spinigera, Hyd. chalcogaster, F. canicularis, and Boettcherisca highlandica Kurahashi & Tan represented by both adults and larvae, whereas C. nigripes, Chrysomya thanomthini Kurahashi & Tumrasvin, M. domestica, Atherigona sp., Parasarcophaga albiceps Meigen, P. taenionota, Sepsidae, Phoridae, and Millichidae were represented by adults only. Faunal succession followed the sequence of dominant flies, i.e., Calliphoridae, Sarcophagidae, Muscidae, Sepsidae, and lastly Stratiomyidae for jungle, or Sepsidae for rural and highland studies. Mites, from suborders Mesostigmata, Prostigmata, Astigmatina, and Oribatida, were also recovered throughout decomposition, which could be used for future implementation in forensic investigations. The data obtained from this study could provide more accurate indicators for local forensic scientists in solving criminal cases especially on the determination of time and primary location of death.

Sepsid flies (Diptera: Sepsidae) are important model insects for sexual selection research. In order to develop mitochondrial (mt) genome data for this significant group, we sequenced the first complete mt genome of the sepsid fly Nemopoda mamaevi Ozerov, 1997. The circular 15,878 bp mt genome is typical of Diptera, containing all 37 genes usually present in bilaterian animals. We discovered inaccurate annotations of fly mt genomes previously deposited on GenBank and thus re-annotated all published mt genomes of Cyclorrhapha. These re-annotations were based on comparative analysis of homologous genes, and provide a statistical analysis of start and stop codon positions. We further detected two 18 bp of conserved intergenic sequences from tRNAGlu-tRNAPhe and ND1-tRNASer(UCN) across Cyclorrhapha, which are the mtTERM binding site motifs. Additionally, we compared automated annotation software MITOS with hand annotation method. Phylogenetic trees based on the mt genome data from Cyclorrhapha were inferred by Maximum-likelihood and Bayesian methods, strongly supported a close relationship between Sepsidae and the Tephritoidea.

Cadaverous entomofauna successions vary according to the region, environment, and climate, and such differences may occur within the same country due to seasonal variations. The present study aimed to analyze and compare the dipterofauna that visit or colonize carcasses in the urban and coastal areas of São Paulo, Brazil, during summer and winter seasons. Four swine (Sus scrofaLinné, 1758) carcasses of approximately 12 kg were used. The animals were previously euthanized and then placed in metal cages covered with a flight intercept trap (Shannon, modified). In total, 10,495 flies from 39 families were collected, with 15 species belonging to the Calliphoridae family, 14 species belonging to the Fanniidae family, 43 species belonging to the Muscidae family, and 22 species belonging to the Sarcophagidae family. Flies from these four families visited all carcasses; however, they did not show the highest visitation frequencies in all of the trials. Species variations occurred between the experiments that were performed at different locations and in different seasons. Furthermore, difference in the number of insects attracted to each stage of decomposition was observed. In addition to the four families highlighted above, the families Phoridae, Sepsidae, Otitidae, and Piophilidae were observed in all carcasses.

Specimens of the enigmatic, monotypic European genus Zuskamira Pont, 1987 (Sepsidae) were initially collected only from the lower central Swedish provinces of Darlana, Uppland and Västmanland. However, the same species was subsequently found much more south in Lower-Saxony and Schleswig-Holstein although Germany is overall well sampled for sepsid flies. Here we report a further (longitudinal) range expansion based on new localities in Southern Finland. New localities for Finland and Sweden are here added and we discuss briefly the habitat requirements of the species.

Die nachfolgenden Arten sind aus der Schweiz gemeldet, müssen jedoch aus der Liste gestrichen werden, da ihr Status ungeklärt ist (nomina dubia), Published as part of Bächli, Gerhard, Merz, Bernhard & Haenni, Jean-Paul, 2014, Dritter Nachtrag zur Checkliste der Diptera der Schweiz, pp. 119-140 in Entomo Helvetica 7 on page 133, DOI: 10.5169/seals-985967, http://zenodo.org/record/8096745

Li, Xuankun, Yang, Ding (2014): Mucha Ozerov and Ortalischema Frey newly recorded from China with descriptions of two new species (Diptera, Sepsidae). Zootaxa 3815 (2): 249-262, DOI: http://dx.doi.org/10.11646/zootaxa.3815.2.5

Key to species of Mucha 1 Arista plumose only on dorsal side (Fig. 5 E)............................................................... 2 - Arista plumose on both sides............................................................. M. plumosa Ozerov 2 Fore tibia straight, ventrally not concave at middle (Figs. 2 A, 6 A).............................................. 3 - Fore tibia ventrally concave at middle (Figs. 1 A, 1 B, 3 A, 5 A, 7 A).............................. M. tzokotucha Ozerov 3 Surstylus pointed ventrally............................................................. M. rectotibialis Iwasa - Surstylus pointed posteroventrally (Figs. 1 D, 3 C, 5 C, 7 C)..................................................... 4 4 Antenna almost yellow; anepisternum mostly glossy, but posteriorly pruinose; hind tibia dark brown, but yellow at apical 1 / 2 (Figs. 1 A, B); lateral view of surstylus short, shorter than length of dorsal epandrium (Figs. 1 D, 5 C)...... M. liangi sp. nov. - Antenna yellowish brown, but ventrally darkened; anepisternum wholly glossy; hind tibia blackish brown with yellow apical ring (Fig. 3 A); lateral view of surstylus long, longer than length of dorsal epandrium (Figs. 3 C, 7 C). M. yunnanensis sp. nov., Published as part of Li, Xuankun & Yang, Ding, 2014, Mucha Ozerov and Ortalischema Frey newly recorded from China with descriptions of two new species (Diptera, Sepsidae), pp. 249-262 in Zootaxa 3815 (2) on page 251, DOI: 10.11646/zootaxa.3815.2.5, http://zenodo.org/record/226959

Sepsis nigripes Meigen, 1826. (Merz 2012c)., Published as part of Bächli, Gerhard, Merz, Bernhard & Haenni, Jean-Paul, 2014, Dritter Nachtrag zur Checkliste der Diptera der Schweiz, pp. 119-140 in Entomo Helvetica 7 on page 127, DOI: 10.5169/seals-985967, http://zenodo.org/record/8096745, {"references":["Merz B. 2012 c. 8.36. 28. Sciomyzoidea, pp. 395 - 397. In: Merz B. (ed.), Liste annotee des Insectes (Insecta) du Canton de Geneve. Instrumenta Biodiversitatis 8: 1 - 532."]}

Sepsis setulosa (Duda, 1926). (Merz 2012c)., Published as part of Bächli, Gerhard, Merz, Bernhard & Haenni, Jean-Paul, 2014, Dritter Nachtrag zur Checkliste der Diptera der Schweiz, pp. 119-140 in Entomo Helvetica 7 on page 127, DOI: 10.5169/seals-985967, http://zenodo.org/record/8096745, {"references":["Merz B. 2012 c. 8.36. 28. Sciomyzoidea, pp. 395 - 397. In: Merz B. (ed.), Liste annotee des Insectes (Insecta) du Canton de Geneve. Instrumenta Biodiversitatis 8: 1 - 532."]}

Key to the Chinese genera of Sepsidae 1 Thorax and abdomen dully shining; mid femur dorsally concaved at middle....................... Toxopoda Macquart - Thorax and abdomen shining; mid femur not concaved at middle............................................... 2 2 Abdomen not constricted between tergites 2 and 3............................................................ 3 - Abdomen constricted between tergites 2 and 3............................................................... 5 3 Br and bm cell united........................................................... Saltella Robineau ���Desvoidy - Br and bm cell separated............................................................................... 4 4 Setae on head reduced, with only 1 vti; fore tarsus flattened and banded black and white............... Ortalishema Fery - Setae on head developed, with 1 oc, 1 pvt, 1 vti, 1 vte, and 1 or; fore tarsus without above structure and colour.............................................................................................. Decachaetophora Duda 5 Supra���alar seta absent; anepisternal seta vestigial or absent............................. Themira Robineau ���Desvoidy - Supra���alar seta present; anepisternal seta present............................................................ 6 6 Arista with plumose or with short rays.................................................................... 7 - Arista bare.......................................................................................... 8 7 Face with well-developed facial carina; flagellomere I strongly lengthened, length nearly 2.5 times longer than width; arista plumose, rays distinctively longer than greatest diameter of arista (Fig. 5 E)............................ Mucha Ozerov - Face without well-developed facial carina; flagellomere I not lengthened, length less than 2 times longer than width; arista with very short rays, longest rays not longer than greatest diameter of arista........................... Aristina Ozerov 8 Fronto���orbital setae developed............................................................ Meroplius Rondani - Front���orbital setae vestigial or absent..................................................................... 9 9 Male fore femur without any spines, setae, and tubercles........................................ Perochaeta Duda - Male fore femur with complex arrangement of spines, setae or tubercles........................................ 10 10 Male fore femur without spines or tubercles, only with a series of very short anteroventral spinules on apical half and a row of well���spaced posteroventral setae; sternite 4 heavily modified, with two pairs of moveable brushy appendages........................................................................................ Nemopoda Robineau ���Desvoidy - Male fore femur not as above; sternite 4 normal, not modified................................................ 11 11 Abdomen in both sexes without distinct striking setae; male surstylus bifurcate at tip................ Dicranosepsis Duda - Abdomen in male and often in female also with distinct striking setae; male surstylus not bifurcate................... 12 12 Middle part of fore femur ventrally with 3���4 strong spines on a strong tubercle, followed by small tubercle frontward bearing 2 pv setae; wing hyaline, without dark spot near apex; epandrium weakly sclerotized.................. Allosepsis Ozerov - Fore femur not as above; wing sometimes with dark spot near apex; epandrium often strongly sclerotized...... Sepsis Fall��n, Published as part of Li, Xuankun & Yang, Ding, 2014, Mucha Ozerov and Ortalischema Frey newly recorded from China with descriptions of two new species (Diptera, Sepsidae), pp. 249-262 in Zootaxa 3815 (2) on page 250, DOI: 10.11646/zootaxa.3815.2.5, http://zenodo.org/record/226959

Genus Ortalischema Frey, 1925 Ortalischema Frey, 1925: 75. Type-species: Sepsis albitarsis Zetterstedt, 1847, by original designation. Protothemira Duda, 1926 a: 25, 26. Type-species: Sepsis albitarsis Zettersted, 1847, by monotypy. Diagnosis. Only 1 vti present; or, oc, pvt, vte absent. Thorax wholly black and thinly pruinose. 2 npl, 1 dc, 1 sa, 1 pa, 1 b sc, 1 sc; pprn absent. Fore tarsus with unique structure and colour. Epandrium with a small rounded anterior lobe and a large oval posterior lobe, both lobes with long thick marginal setulae., Published as part of Li, Xuankun & Yang, Ding, 2014, Mucha Ozerov and Ortalischema Frey newly recorded from China with descriptions of two new species (Diptera, Sepsidae), pp. 249-262 in Zootaxa 3815 (2) on page 256, DOI: 10.11646/zootaxa.3815.2.5, http://zenodo.org/record/226959, {"references":["Frey, R. (1925) Zur Systematik der Diptera Haplostomata II. Fam. Sepsidae. Notulae Entomologicae, 5, 69 - 76."]}

Genus Mucha Ozerov, 1992 Mucha Ozerov, 1992: 147. Type-species: Mucha tzokotucha Ozerov, 1992, by original designation. Diagnosis. Postgena wide; face with well-developed facial carina between antennae. Flagellomere elongated; arista plumose (mostly on dorsal side, except Mucha plumosa Ozerov with short rays on ventral side of arista). 1 oc, 1 vti, 1 vte, 1 pvt, or vestigial; anal vein 8 times as long as width of bm cell; alula not developed, with microtrichiae marginally. Abdominal syntergite 1 + 2 with one pair of strong setae., Published as part of Li, Xuankun & Yang, Ding, 2014, Mucha Ozerov and Ortalischema Frey newly recorded from China with descriptions of two new species (Diptera, Sepsidae), pp. 249-262 in Zootaxa 3815 (2) on page 250, DOI: 10.11646/zootaxa.3815.2.5, http://zenodo.org/record/226959, {"references":["Ozerov, A. L. (1992) Novyy rod i tri novykh vida murav'evidok (Diptera, Sepsidae) iz V'etnama [New genus and three new species of sepsid flies (Diptera, Sepsidae) from Vietnam]. Zoologicheskiy zhurnal, 73 (3), 147 - 150."]}

3. Mucha yunnanensis sp. nov. (Figs. 3; 7) Diagnosis. Vibrissal angle with 3 strong setae. Antenna yellowish brown, but ventrally darkened. Thoracic anepisternum wholly glossy. Hind tibia blackish brown with yellow apical ring (Fig. 3 A,). Lateral view of surstylus long, de: as: ps = 1: 1.3: 1.5 (Figs. 3 C, 7 C). Description. Male. Body length 4.5 mm, wing length 3.2 mm. Head mostly black; frons shiny and blackish brown apically; face almost brown except facial carina yellow; facial orbit black to blackish brown; gena yellow; face and occiput thinly pruinose. 1 oc, 1 vti, 1 vte, 1 pvt, or vestigial. Vibrissal angle with 3 strong setae; genal margin with 8 peristomal setae in a row, and with 1 strong seta at lower margin near postgena. Antenna yellowish brown, but ventrally darkened; first flagellomere distinctively elongated, 2.4 times longer than wide, pubescent; arista brown, long plumose on dorsal side. Proboscis yellow with brown setulae; palpus yellow except apex yellowish brown, with black setulae. Thorax mostly blackish brown, except protonotopleural lobe yellowish brown ventrally, propisternum yellowish brown, proepimeron and antepisternum brown. Thorax mostly pruinose, except anepisternum wholly glossy and katepisternum shining anteriorly. 1 pprn, 2 npl, 2 dc, 1 sa, 1 pa, 1 m, 1 b sc, 1 sc. Legs mostly blackish brown, except fore and mid coxae brown with blackish brown vertical band; hind femur yellow at base; fore tibia entirely yellow, mid tibia yellow at apical 1 / 2, hind tibia with yellow apical ring, and all tarsomeres 1-3 yellow. Fore femur ventrally with group of 3 pv setae, 1 v seta, and 1 weak av seta near middle, followed by small tubercle frontward bearing 2 pv setae, and with 7 ventral setulae on distal 1 / 3; mid and hind femora without any distinct setae or spines (Figs. 3 A, 7 A, 7 B). Wing hyaline, slightly tinged brown, without dark spot near apex; veins dark brown; anal vein 8 times as long as width of bm cell; alula not developed, about 1.5 times as wide as bm cell, with microtrichiae marginally. Halter white, but blackish brown basally. Abdomen mostly blackish brown and shiny, except syntergite 1 + 2 brown, tergites 3-5 and syntergite 7 + 8 brownish yellow laterally and apically. Syntergite 1 + 2 to tergite 5, and syntergite 7 + 8 normal, tergite 6 missing. Syntergite 1 + 2 with one pair of striking setae; tergites 3���5 each with pair of marginal and dorsal setulae. Sternite 3 longitudinally elongated, sternite 4 oval, each with setulae (Fig. 3 B). Epandrium with one pair of strong dorsal setae and 3 pairs of posterior setae. Surstylus apically tapering, not bifurcate. Relative length ratio of epandrium and surstylus: de: ade: ave: as = 1: 2.25: 3.25: 1.3; de: pde: pve: ps = 1: 2.1: 2.4: 1.5 (Figs. 3 C, 7 C). Female. Unknown. Type material. HOLOTYPE ♂, CHINA, Yunnan, Baoshan, Baihualing, 1575 m, 2013. VII. 16, Xuankun Li. Distribution. Oriental: Yunnan. Etymology. This specific epithet refers to the type locality Yunnan. Remarks. This species is closely similar to M. rectotibialis Iwasa, but is distinguishable from it by the shape of male surstylus. It is also similar to M. liangi sp. nov., but can be separated from it by the following features: antenna yellowish brown, but ventrally darkened; thoracic anepisternum wholly glossy; hind tibia blackish brown with yellow apical ring; lateral view of surstylus long, de: as: ps = 1: 1.3: 1.5., Published as part of Li, Xuankun & Yang, Ding, 2014, Mucha Ozerov and Ortalischema Frey newly recorded from China with descriptions of two new species (Diptera, Sepsidae), pp. 249-262 in Zootaxa 3815 (2) on pages 254-256, DOI: 10.11646/zootaxa.3815.2.5, http://zenodo.org/record/226959

1. Mucha liangi sp. nov. (Figs. 1, 5) Diagnosis. Head mostly black with blue metal luster. Vibrissal angle with 3 strong setae. Antenna almost yellow. Thoracic anepisternum mostly glossy, but posteriorly pruinose. Hind tibia yellow at apical 1 / 2 (Figs. 1 A, 1 B). Lateral view of surstylus short, de: as: ps = 1: 0.57: 0.57 (Figs. 1 D, 5 C). Description. Male. Body length 4.2 mm, wing length 3.1 mm. Head mostly black with blue metal luster; frons shiny and blackish brown apically; face almost brown except facial carina yellow; facial orbit black to blackish brown; gena yellow, occiput thinly pruinose. 1 oc, 1 vti, 1 vte, 1 pvt, or vestigial. Vibrissal angle with 3 strong setae; genal margin with 8 peristomal setae in a row, and with 1 strong seta at lower margin near postgena. Antenna almost yellow; first flagellomere distinctively elongated, 2.4 times longer than wide, pubescent; arista yellowish brown, long plumose on dorsal side. Proboscis yellow with brown setulae; palpus yellow except apex yellowish brown, with black setulae. Thorax mostly blackish brown, except protonotopleural lobe yellowish brown ventrally, and propisternum yellowish brown. Thorax mostly pruinose, except anepisternum mostly glossy but posteriorly pruinose; katepisternum shining anteriorly. 1 pprn, 2 npl, 2 dc, 1 sa, 1 pa, 1 m, 1 b sc, 1 sc. Legs mostly dark brown, except fore and mid coxae brown with blackish brown vertical band; hind femur yellow at base; fore tibia entirely yellow, mid and hind tibiae yellow at apical 1 / 2; all tarsomeres 1���3 yellow. Fore femur ventrally with group of 3 pv setae, 1 v seta, and 1 av seta near middle, followed by small tubercle frontward bearing 2 pv setae, and with 5 ventral setulae on distal 1 / 3; mid and hind femora without any distinct setae or spines (Figs. 1 A, 5 A, 5 B). Wing hyaline, slightly tinged brown, without dark spot near apex; veins dark brown; anal vein 8 times as long as width of bm cell; alula not developed, about 1.5 times as wide as bm cell, with microtrichiae marginally. Halter white, but blackish brown basally. Abdomen mostly blackish brown and shiny, except syntergite 1 + 2 brown, tergite 3 brown laterally, tergites 4-5 and syntergite 7 + 8 brownish yellow apically. Syntergite 1 + 2 to tergite 5, and syntergite 7 + 8 normal, tergite 6 missing. Syntergite 1 + 2 with one pair of striking setae; tergites 3-5 each with pair of marginal and dorsal setulae. Sternite 3 longitudinally elongated, sternite 4 oval, each with lateral and apical setulae (Fig. 1 C). Epandrium with one pair of strong dorsal setae and 3 pairs of posterior setae. Surstylus apically tapering, not bifurcate. Relative length ratio of epandrium and surstylus: de: ade: ave: as = 1: 1.71: 2.57: 0.57; de: pde: pve: ps = 1: 1.43: 2.07: 0.57 (Figs. 1 D, 5 C). Female. Body length 5.0 mm, wing length 3.4 mm. Similar to male, but fore femur, without above specialized features (Fig. 1 B). Type material. HOLOTYPE ♂, CHINA, Guangxi, Baise, Cenwanglaoshan Mt., Dalongping, 1300 m, 2013. VII. 29, Feiyang Liang. PARATYPE: 1 ♀, same data as holotype. Distribution. Oriental: Guangxi. Etymology. This species is named after the type collector Mr. Feiyang Liang. Remarks. This species is very similar to M. yunnanensis sp. nov., but can be separated from it by the following features: antenna almost yellow; thoracic anepisternum mostly glossy, but posteriorly pruinose; fore femur with 5 ventral setae on distal 1 / 3; hind tibia yellow at apical 1 / 2; lateral view of surstylus short, de: as: ps = 1: 0.57: 0.57., Published as part of Li, Xuankun & Yang, Ding, 2014, Mucha Ozerov and Ortalischema Frey newly recorded from China with descriptions of two new species (Diptera, Sepsidae), pp. 249-262 in Zootaxa 3815 (2) on page 251, DOI: 10.11646/zootaxa.3815.2.5, http://zenodo.org/record/226959

2. Mucha tzokotucha Ozerov (Figs. 2, 6) Mucha tzokotucha Ozerov, 1992 c: 148 (Mucha). Type-locality: Ba Vi 70 km NW Hanoi (VIETNAM: Hanoi); Holotype ♂, in ZISP. Redescription. Male. Body length 4.5 mm, wing length 3.2 mm. Head mostly black; frons shiny and blackish brown apically; face almost brown; facial orbit black to blackish brown; gena yellowish brown; face and occiput thinly pruinose. 1 oc, 1 vti, 1 vte, 1 pvt, or vestigial. Vibrissal angle with 1 strong and 1 weak setae; genal margin with 7 peristomal setae in a row, and with 1 strong seta at lower margin near postgena. Antenna almost yellow; first flagellomere distinctively elongated, 2.4 times longer than wide, pubescent; arista brown, long plumose on dorsal side. Proboscis yellow with brown setulae; palpus yellow except apex yellowish brown, with black setulae. Thorax mostly blackish brown, except protonotopleural lobe yellowish brown ventrally and propisternum yellowish brown. Thorax mostly pruinose, except anepisternum glossy and katepisternum shining anteriorly. 1 pprn, 2 npl, 2 dc, 1 sa, 1 pa, 1 m, 1 b sc, 1 sc. Legs mostly blackish brown, except fore and mid coxae brown with blackish brown vertical band; hind femur yellow at base; fore tibia entirely yellow, mid and hind tibiae yellow at apical 1 / 2, and all tarsomeres 1-3 yellow. Fore femur ventrally with group of 3 pv setae, 1 v seta, and 1 weak av seta near middle, followed by small tubercle frontward bearing 2 pv setae, and with 7 ventral setulae on distal 1 / 3; mid and hind femora without any distinct setae or spines; fore tibia ventrally concave at middle (Figs. 2 A, 6 A, 6 B). Wing hyaline, slightly tinged brown, without dark spot near apex; veins dark brown; anal vein 8 times as long as width of bm cell; alula not developed, about 1.5 times as wide as bm cell, with microtrichiae marginally. Halter white, but blackish brown basally. Abdomen mostly brown and shiny, except syntergite 1 + 2, tergites 3-5 and syntergite 7 + 8 brownish yellow laterally and apically. Syntergite 1 + 2 to tergite 5, and syntergite 7 + 8 normal, tergite 6 missing. Syntergite 1 + 2 with one pair of striking setae; tergites 3-5 each with pair of marginal and dorsal setulae. Sternite 3 longitudinally elongated, sternite 4 oval, each with setulae (Fig. 2 B). Epandrium with one pair of strong dorsal setae and 3 pairs of posterior setae. Surstylus apically tapering, not bifurcate. Relative length ratio of epandrium and surstylus: de: ade: ave: as = 1: 1.74: 2.83: 1.30; de: pde: pve: ps = 1: 1.83: 2.22: 1.17 (Figs. 2 C, 6 C). Female. Unknown. Speciments examined. 1 ♂ China, Jangxi, Jinggangshan, Jingzhushan Mt., 600 m, 2012. X. 10, Ding Yang. Distribution. Oriental: China (Jiangxi), Vietnam. New to China. Remarks. This species is recorded from China for the first time. This species can be separated from other species of Mucha by the following features: vibrissal angle with 1 strong and 1 weak setae; fore tibia ventrally concave at middle., Published as part of Li, Xuankun & Yang, Ding, 2014, Mucha Ozerov and Ortalischema Frey newly recorded from China with descriptions of two new species (Diptera, Sepsidae), pp. 249-262 in Zootaxa 3815 (2) on pages 253-254, DOI: 10.11646/zootaxa.3815.2.5, http://zenodo.org/record/226959, {"references":["Ozerov, A. L. (1992) Novyy rod i tri novykh vida murav'evidok (Diptera, Sepsidae) iz V'etnama [New genus and three new species of sepsid flies (Diptera, Sepsidae) from Vietnam]. Zoologicheskiy zhurnal, 73 (3), 147 - 150."]}

4. Ortalischema maritima Ozerov (Figs. 4, 8) Ortalischema maritima Ozerov, 1985: 1268. Type-locality: "����йхэ" ["Mayche", now = Artemovka] (RUSSIA: Primorskiy Kray); Holotype ♂, in ZISP. Redescription. Male. Body length 4.5 mm; wing length 3.5 mm. Head roundish, mostly black; frons dark brown apically; face light brown; facial orbit brownish yellow; gena and occiput shiny and thinly pruinose. Only 1 vti present; or, oc, pvt, vte absent. Setae on vibrissal angle vestigial; genal margin with at least 10 peristomal setae in a row and 1 strong seta at genal angle. Antennal dark brown; first flagellomere pubescent; arista blackish brown except base yellow. Proboscis black with blackish brown setulae; palpus black with black setulae. Thorax wholly black with thin pruinosity, except greater ampulla and anemiperon anteriorly shining. 2 npl, 1 dc, 1 sa, 1 pa, 1 b sc, 1 sc; pprn absent. Legs mostly black, except mid and hind femora dark brown at base, fore and mid tibiae dark brown at both ends; fore tarsomere 1 white, tarsomere 2 white except apex black, mid tarsomere 1 dark brown. Fore femur with two rows of short spines anteroventrally and posteroventrally; fore tarsomeres 1-3 thickened; mid tarsus with long thick ventral setulae. Setae and setulae on legs black (Fig. 4 A). Wing hyaline, slightly tinged brown, without dark spot near apex; veins dark brown. Alula white with marginal microtrichiae. Halter pale yellow, black basally. Abdomen black and shiny. Tergites sparsely clothed with setulae, without striking setae. Epandrium broad in lateral view, covered with setae, marginal setae longer than others; epandrium with a small rounded anterior lobe and a large oval posterior lobe, both lobes with long thick marginal setulae. Surstylus small, with some setae (Figs. 8 A, 8 B). Female. Body length 4.7 mm; wing length 3.6 mm. Very similar to male, except fore tarsomere 1 black, tarsomere 2 white, tarsomere 3 white with brown apex; mid tarsomere 1 black without long thick ventral setulae (Fig. 4 B). Material examed. China: 3 ♂, 2 ♀, Ningxia, Liupanshan, Liangdianxia, 2000 m, 2007. VI. 28, Gang Yao; 7 ♂, 2 ♀, Ningxia, Liupanshan, Liangdianxia, 2000 m, 2007. VI. 28, Qibiao Dong; 1 ♂, Ningxia, Liupanshan, Liangdianxia, 2000 m, 2007. VI. 29, Gang Yao; 4 ♂, 2 ♀, Ningxia, Liupanshan, Liangdianxia, 2000 m, 2007. VI. 29, Qibiao Dong; 10 ♂, Ningxia, Liupanshan, Liangdianxia, 2000 m, 2007. VI. 30, Gang Yao; 4 ♂, 2 ♀, Ningxia, Liupanshan, Liangdianxia, 2000 m, 2007. VI. 30, Qibiao Dong. Distribution. China, Japan (Hokkaido I.), Mongolia, Russia (Primorskiy Kray, Sakhalinskaya Oblast���). New to China. Remarks. This species is recorded from China for the first time. Iwasa (1985) described the species with detailed illustrations, but had misidentified it as O. albitarsis. Iwasa (1995) corrected this and added illustrations of both species., Published as part of Li, Xuankun & Yang, Ding, 2014, Mucha Ozerov and Ortalischema Frey newly recorded from China with descriptions of two new species (Diptera, Sepsidae), pp. 249-262 in Zootaxa 3815 (2) on pages 256-257, DOI: 10.11646/zootaxa.3815.2.5, http://zenodo.org/record/226959, {"references":["Ozerov, A. L. (1985) K taksonomii roda Ortalischema (Diptera, Sepsidae) [To the taxonomy of the genus Ortalischema (Diptera, Sepsidae)]. Zoologicheskiy zhurnal, 64 (8), 1267 - 1269.","Iwasa, M. (1985) Supplementary notes on the Sepsidae from Japan (Diptera). Kontyu, 53 (4), 632 - 638.","Iwasa, M. (1995) Revisional Notes on the Japanese Sepsidae (Diptera). Japanese Journal of Sanitary Zoology, 63 (4), 781 - 797."]}

Les caracteres impliques dans les interactions sexuelles entre les mâles et les femelles ont tendance a diverger plus rapidement que les autres caracteres. Plusieurs hypotheses cherchent a expliquer ce patron evolutif. L'examen de l'appariement de deux appareils mâles specifiques aux structures femelles durant la cour et l'accouplement chez la mouche Sepsis cynipsea (L., 1758) a permis de verifier les predictions de deux de ces hypotheses, celle de la serrure et de la cle et celle de la coevolution morphologique sexuelle antagoniste. Contrairement aux predictions des deux hypotheses, les femelles ne possedent pas de modifications des structures (bases des ailes, 6 e sternite abdominal) qui sont en interaction avec les modifications specifiques des pattes anterieures des mâles, ni avec celles de leurs surstyles genitaux. Les mâles endommagent de petites plages de microtriches sur les ailes des femelles avec leurs pattes; la morphologie de la femelle ne montre, cependant, aucune signe des adaptations de l'aile et de l'abdomen prevues par la coevolution morphologique antagoniste. Les donnees concernant l'hypothese de rechange de la coevolution du comportement sexuel antagoniste chez la femelle en reaction a la morphologie du mâle sont encore moins claires; l'hypothese ne reussit pas a expliquer la compression rythmique soutenue, apparemment stimulatrice, exercee par les genitalias mâles et l'absence de reactions de defense de la part de la femelle a cette compression. Les mouvements des surstyles mâles durant l'accouplement laissent croire qu'ils servent a stimuler la femelle. La base de l'aile de la femelle possede, semble-t-il, des organes sensoriels pres des surfaces palpees par le mâle, tel que prevu par l'hypothese de rechange traditionnelle du choix de la femelle pour expliquer l'evolution divergente rapide. Les surstyles genitaux du mâle servent aussi a des interactions precopulatrices inedites: une paire de processus, encore non decrits, a la base retiennent probablement la femelle et la compriment aussi sans doute rythmiquement durant la copulation.

In this study we use sensitivity analysis sensuWheeler (1995) for a matrix entirely composed of DNA sequences. We propose that not only congruence but also phylogenetic structure, as measured by character resampling, should be used to choose among competing weighting regimes. An extensive analysis of a five-gene data set for Themira (Sepsidae: Diptera) reveals that even with different ways of partitioning the data, measures of topological congruence, character incongruence, and phylogenetic structure favor similar weighting regimes involving the down-weighting of transitions. We furthermore use sensitivity analysis for obtaining empirical evidence that allows us to select weights for third positions, deciding between treating indels as fifth character states or missing values, and choosing between manual and computational alignments. For our data, sensitivity analysis favors manual alignment over a Clustal-generated numerical alignment, the treatment of indels as fifth character states over considering them missing values, and equal weights for all positions in protein-encoding genes over the down-weighting of third positions. Among the topological congruence measures compared, symmetric tree distance performed best. Partitioned Bremer Support analysis reveals that COI contributes the largest amount of support for our phylogenetic tree for Themira. © The Willi Hennig Society 2005.

Previous univariate studies of the fly Sepsis cynipsea (Diptera: Sepsidae) have demonstrated spatiotemporally variable and consequently overall weak sexual selection favouring large male size, which is nevertheless stronger on average than fecundity selection favouring larger females. To identify specific target(s) of selection on body size and additional traits possibly affecting mating success, two multivariate field studies of sexual selection were conducted. In one study using seasonal replicates from three populations, we assessed 15 morphological traits. No clear targets of sexual selection on male size could be detected, perhaps because spatiotemporal variation in selection was again strong. In particular, there was no (current) selection on male abdomen length or fore coxa length, the only traits for which S. cynipsea males are not smaller than females. Interestingly, copulating males had a consistently shorter fore femur base, a secondary sexual trait, and a wider clasper (hypopygium) gap, an external genital trait. In a second study using daily and seasonal replicates from one population, we included physiological measures of energy reserves (lipids, glucose, glycogen), in addition to hind tibia length and fluctuating asymmetry (FA) of all pairs of legs. This study again confirmed the mating advantage of large males, and additionally suggests independent positive influences of lipids (the long-term energy stores), with effects of glucose and glycogen (the short-term energy stores) tending to be negative. FA of paired traits was not associated with male mating success. Our study suggests that inclusion of physiological measures and genital traits in phenomenological studies of selection, which is rare, would be fruitful in other species.

Mating more than once is extremely costly for females in many species, making the near ubiquity of polyandry difficult to understand. However, evidence of mating costs for males is much rarer. We investigated the effects of copulation on longevity of male and female flies (Saltella sphondylli). We also scrutinized potential fecundity and fertility benefits to females with differing mating history. Copulation per se was found to decrease the longevity of males but not that of females. However, when females were allowed to lay eggs, females that mated died earlier than virgin females, indicating costs of egg production and/or oviposition. Thus, although longevity costs of copulation are higher for males, reproduction is nevertheless costly for females. We also found no differences in fecundity or fertility relative to female mating history. Results suggest that polyandry may be driven by minor costs rather than by major benefits in this species.

Probable ancestral types of courtship behavior for most sepsid flies were deduced from the behavior and morphology of a species in the relatively basal genus Themira. At least three of the five behavior patterns used by Themira minor (Haliday) males to court females prior to and during copulation are shared with other, more derived groups. During copulation the male's sternal lobes were rotated so that their brushes of long setae were fanned apart and may tap the female. The morphology of the tips of the male's genitalic surstyli and the muscles attached to them indicate that they are also moveable. This trait, which occurs in several other distantly related species, probably serves to squeeze the female during copulation. Thus, contrary to the impression from previous studies, elaborate courtship before and during copulation may be widespread among sepsids.

Este artículo intenta organizar algunas ideas teóricas sobre posibles conflictos entre machos y hembras en cuanto a la cópula. Las ideas se ilustran con datos nuevos de una familia de moscas (Sepsidae), y con otros datos ya publicados sobre otros insectos. El hecho de que una hembra opone resistencia a la penetración por el macho no implica en sí mismo que la cópula es forzada, ya que la resistencia de la hembra también puede funcionar como una prueba del macho, para así tamizar entre diferentes machos y conseguir hijos superiores. Se contrastan dos hipótesis para el macho, la de la cópula por coerción vs. la de la cópula por persuasión; también se contrastan, desde la perspectiva de la hembra, dos hipóteses relacionadas, la resistencia para evitar a todo macho en forma no selectiva vs. la resistencia para tamizar entre machos. Cuando la morfología de la genitalia de los dos sexos es tal que el macho no es físicamente capaz de forzar la intromisión, como es el caso en las moscas sépsidas y probablemente en muchos otros insectos (a pesar de algunas publicaciones que sugieren lo contrario), las hipótesis contrastantes para cada sexo pueden resolverse con datos sobre el contexto ecológico en el cual ocurre la interacción macho-hembra. Se realizó un análisis de esta clase con los sépsidos. Parece más probable que la resistencia energética que la hembra pone al macho sea un esfuerzo para tamizar entre machos que un esfuerzo para evitar la cópula en forma no selectivaFemale resistance behavior that occurs prior to intromission does not by itself imply forced copulation. Such behavior may function instead as a test of the male in order to favor some males over others, or to induce the male to desist. Thus, male persistence and forcefulness may sometimes be better described as persuasion rather than coercion. Under the persuasion hypothesis, the male only gains intromission due to an active response of the female. Under the coercion hypothesis, male and female are opposed in a physical battle which the female loses if copulation occurs. In species in which males are morphologically incapable of forcing intromission without active female cooperation (I argue here that this is probably a very common situation), data on the behavioral and ecological context in which resistance occurs can distinguish between the two possibilities. Partially congruent functions of resistance, seen from the female point of view, are female resistance to screen (male persuasion), and female resistance to avoid males non-selectively (male coercion). Sepsid flies illustrate these ideas. Females often struggle energetically in apparent attempts to dislodge mounted males and to prevent intromission, and males grasp females with powerful species-specific structures on their front legs and genitalia. This suggests the possibility of coerced intromission. But behavioral and morphological evidence demonstrate that active female cooperation occurs at the moment of intromission, and that males are probably dependent on this cooperation because they are not morphologically equipped to force their genitalia into those of an uncooperative female. Despite the impression from previous publications, male insects in general may seldom be able to achieve intromission by genitalic force. The species-specific forms of the grasping genitalia of male sepsids are probably not the result of an evolutionary arms race between coercive males and unselectively resistant females

Four hypotheses that could explain the elaborate species-specific morphology of the clasping organs on the front legs of male Archisepsis diversiformis flies were tested: direct male–male combat, mechanical fit, male–female conflict of interests, and male stimulation of the female. Experimental modification of the shape of the male clasper and of the female wing base where the male clasped the female both strongly reduced the chances that a mount would result in copulation. This reduction was not predicted by the male–male combat hypothesis but was predicted by the others. Males in the field did nave to fight other males to remain mounted on females, as expected by the male–male combat hypothesis. Reduced male copulatory success was not due to inferior male ability to grasp and hold onto the female's wings, as predicted by the mechanical fit and male–female conflict hypotheses but to a reduction in the likelihood that the female would allow intromission, as predicted by the stimulation hypothesis. By a process of elimination, and in combination with data from a previous morphological study, the data support the hypothesis that the species-specific aspects of grasping organs in these flies function to stimulate females. Further behavioral data will be needed to test alternative possibilities.

Finding, assessing, rejecting, and copulating with a mate is assumed to carry fitness costs, particularly for females, that have to be traded off against fitness benefits of mating such as increased fecundity, fertility, longevity, or better quality offspring. Female dung flies, Sepsis cynipsea (Diptera: Sepsidae), typically attempt to dislodge mounted males harassing them by vigorous shaking. Shaking duration has been shown to reflect both direct and indirect female choice in this species. The latter is an expression of the females' general reluctance to mate due to presumed costs of mating. We investigated the costs of copulation in the laboratory. Females were randomly assigned to one of three treatment groups and allowed to copulate either not at all, once, or twice. The males' armored genitalia injured females internally during copula. Injuries were visible as sclerotized scars in the female ovipositor, and their occurrence increased with mating frequency. Presumably due to these injuries, mated females showed higher mortality. This effect was statistically independent from additional costs of reproduction related to oviposition, as copulation also increased lifetime egg production and tended to augment oviposition rate (eggs per day), while fertility (proportion of offspring emerged) was unaffected. We thus found high mortality costs of copulating, indicating substantial sexual conflict, which helps explain female reluctance to mate in this species. Copyright 2002.

Mating is generally assumed to carry costs, particularly for females, which have to be traded off against each other and against the fitness benefits of mating. To understand any particular mating system and the evolution of sexual conflict, these costs have to be evaluated. Female dung flies, Sepsis cynipsea (Diptera: Sepsidae), typically attempt to dislodge mounted males by vigorous shaking. Such female reluctance to mate can only evolve if the cost of avoiding matings does not exceed the cost of copulation. We investigated female precopulatory costs of assessing and rejecting males in terms of increased predation, wing injuries, and (indirectly) energetics, all ultimately affecting mortality, and compared them to the costs of copulation assessed in this and a companion study. Females housed with a male had lower survivorship than females housed with another female. This was largely due to the costs of copulation rather than presumed energetic costs of avoiding males, which were minor. Male harassment augmented female wing injuries, which accumulate with age in the field and laboratory, but in laboratory experiments using one common predator, wing injuries did not increase the susceptibility of S. cynipsea to predation, nor did their mating behavior per se. Instead, predation was highest and survivorship lowest in all-male groups, probably because males are more active in search of females and harass each other. Overall, the precopulatory costs of mate assessment and rejection were low relative to the costs of copulating, explaining female reluctance behavior in this and possibly other species. Copyright 2002.

Several possible explanations for the elaborate species-specific morphology of male front leg clasping organs were tested by comparing six species of Archisepsis , Palaeosepsis and Microsepsis flies. The only previously published hypothesis regarding these clasping organs was refuted by the finding that species-specific portions of the male femur and tibia consistently meshed tightly with prominent veins and folds in the female's wing, rather than meshing with each other. Female wing morphology in the region grasped by the male was relatively uniform and in general did not vary in ways that would prevent non-conspecific males from grasping them, arguing in all but one species against both simple lock-and-key and male–female conflict of interests hypotheses based on morphology. Interspecific differences in male front leg morphology generally represent alternative ways to accomplish the same basic mechanical function of holding tightly onto the relatively invariant female. Despite the fact that female resistance behaviour indicates that male–female conflict over male mounting is common, only one female wing structure in one species resembled an anti-clasper device, giving a second reason to doubt the morphological male–female conflict of interest hypothesis, at least for five of the six species. The positions of probable sensory structures on the wings of females were relatively similar in different species and did not correspond in any obvious way to species-specific features of male clasping structures. This, plus the intraspecific variation in both the positions of these sensilla and the exact site where the male grasped the female's wing, argued against simple «sensory lock-and-key» ideas about male front leg function. By a process of elimination, it appears that generalized female receptors are able to sense species-specific differences in male front legs. This idea was supported by increased female rejection behaviour in cross-specific pairs.

O presente trabalho teve como objetivo estudar a emergência de Haematobia irritans em fezes de bovinos mantidas a campo e em laboratório, além de observar a presença de outros dípteros associados às massas fecais. Foram utilizados dois grupos de fezes bovinas, um à campo e outro em laboratório, sendo cada grupo formado por cinco bolos fecais frescos de tamanho mediano (cerca de 30 cm de diâmetro), aparência opaca, cor esverdeada, presença de fina crosta externa e consistência firme. As fezes foram cobertas com gaiolas de emergência de formato piramidal, apresentando na parte superior um orifício onde foi acoplado um frasco removível; a substituição das fezes foi realizada quinzenalmente. Foram coletados um total de 355 espécimens de H. irritans, sendo 151 provenientes das gaiolas mantidas no campo e 204 em laboratório. Estas diferenças deveram-se provavelmente à menores oscilações de temperatura verificadas no laboratório do que àquelas ocorridas no campo, além da ação de artrópodes predadores, competidores e parasitóides que, provavelmente, interferiram na emergência de H. irritans nas fezes bovinas mantidas no campo. Foi observado maior número de fêmeas em comparação com machos em ambas as condições investigadas. Além de H. irritans, obteve-se outros dípteros associados às fezes bovinas, pertencentes as seguintes famílias: Aulacigastridae, Muscidae, Psychodidae, Sarcophagidae, Sepsidae, Tachinidae e Ulidiidae. Sepsidae foi a mais abundante, com 5.224 exemplares do total de 8.928 dípteros obtidos, seguida por Sarcophagidae com 2.235 espécimens coletados, Muscidae com 1.357, Aulacigastridae com 54, Psychodidae com 46, Ulidiidae com 6 e Tachinidae com 5 exemplares. This study was carried out in order to investigate the emergence of adult flies of Haematobia irritans in cattle dung maintained in the field and in the laboratory, as well as other flies associated with dung pats. Two groups of cattle dung were used, one in the field and another in the laboratory, each group consisting of five fresh dung pats of medium size (about 30 cm in diameter), opaque appearance, presence of a thin external crust and firm consistency. The dung pats were covered with emergence cages in pyramidal format, presenting a hole in the superior part where a removable flask was coupled, and the substitution of the dung was done every 2 weeks. A total of 355 specimens of H. irritans was collected, 151 of which were obtained in the field and 204 specimens in the laboratory. These differences probably were due to smaller temperature variations in the laboratory than in the field, besides the action of arthropodal predators, competitors and parasites of horn fly pupae that probably interfered in H. irritans emergence in cattle dung under field conditions. A higher number of females were observed in both conditions. Besides H. irritans, there were collected other flies associated with bovine manure belonging to the following families: Aulacigastridae, Muscidae, Psychodidae, Sarcophagidae, Sepsidae, Tachinidae and Ulidiidae. Sepsidae was the most abundant, with 5,224 specimens of the total of 8,928 flies collected, followed by Sarcophagidae with 2,235, Muscidae with 1,357, Aulacigastridae with 54, Psychodidae with 46, Ulidiidae with 6 and Tachinidae with 5 specimens.

We investigated the mechanisms of sexual selection in the common dung fly Sepsis cynipsea and how these affect selection on body size at the population level. Because of the presumed costs associated with mating, we predicted that there would be a decrease in the general reluctance of females to mate with any particular male at higher male densities at the mating site, a fresh cow pat, resulting in indirect female choice and a decrease in the strength of sexual selection. In contrast, classical direct female choice and male-male competition should result in increased selection intensities because more opportunities for choice and competition exist at higher densities. Female reluctance to mate and female assessment of males are expressed in prominent female behaviour to repel mates in several insect species, including S. cynipsea. Laboratory pair-wise choice experiments showed that large males were more likely to obtain copulations, which also ensued more promptly, suggesting female assessment of male quality (direct female choice). There was a basic influence of male activity but little further effect of male scramble competition on the outcome of mating. Another laboratory experiment showed a decrease in female shaking duration per male, associated with an asymptote in the shaking duration per female, as male density and harassment increased, but did not show the increase in mating frequency predicted by the female reluctance hypothesis. A study estimating sexual selection differentials in the field showed that directional selection for larger males was present overall and was negatively related to seasonally mediated variation in male density. Our study suggests that direct female choice in combination with indirect female choice (due to an interaction of female reluctance to mate and male persistence) is most consistent with the behavioural and selection patterns observed in S. cynipsea, but male effects cannot be definitively excluded.

Insect activity was significantly reduced in dung from cattle treated with a recommended topical dose (500 μg/kg) of the endectocide, ivermectin. Reductions affected an ecologically and taxonomically diverse group of insects including coprophagous flies, parasitic wasps, and both predacious and coprophagous beetles. Whereas some species appeared unaffected by faecal residues, the emergence of other species was reduced in dung voided ≤ 12 weeks post-application. The species most affected were the flies, Sepsis sp. and Coproica mitchelli (Malloch). eucoilid wasps. and the beetles, Cercyon quisquilius (Linnaeus) and C. pygmaeus (Illiger). Results were consistent within and between years. Similar reductions have been reported previously, but typically for a much shorter post-application period. The greater duration of effect observed in the current study is attributed to a more accurate bioassay, species-level identifications, and the use of sample sizes generally larger than those used in previous studies. Reduced insect activity was associated with slower dung pat degradation. When ivermectin was added directly to dung, at levels previously reported to occur in dung from treated cattle, the treated dung had not appreciably degraded after 340 days in the field. In contrast, untreated dung pats deposited at the same time and place had largely degraded after 80 days in the field. The effect of ivermectin use on pasture quality is probably ‘moot’ in southern Alberta because of the current pattern of use. Should this pattern change, however, results of the present study validate the need for future studies to assess the off-target effects of ivermectin use on pasture quality.

1. Seasonality is a prime selective factor promoting genetic differentiation of populations. Local adaptation in diapause response was investigated in the two geographically and altitudinally widespread dung flies Scathophaga stercoraria (Diptera: Scathophagidae) and Sepsis cynipsea (Diptera: Sepsidae). 2. Replicate sibships from three sites in Switzerland with low and high altitude dung fly populations were raised in a common laboratory environment simulating the natural decreasing photoperiod and temperature regimen before winter. From field phenologies, the critical photoperiod inducing diapause was predicted to be longer for high than for low altitude populations (12 vs. 10 h for Sc. stercoraria, and 12.25 vs. 11 h for Se. cynipsea) if they are locally adapted. 3. Contrary to expectation and to Sc. stercoraria, which diapauses in the pupal stage, Se. cynipsea diapauses in the adult stage. 4. Low but significant levels of genetic differentiation in pre-winter adult emergence were evident between low and high altitude populations of both species, but they were far from the differences predicted. Scathophaga stercoraria also showed geographical differentiation independent of altitude. 5. Sepsis cynipsea females stopped reproducing at some point before winter, but altitude did not affect the timing of adult diapause. High altitude females and females that did not initiate reproduction before winter survived the simulated winter better. 6. Both species largely used temperature rather than photoperiod as a cue inducing winter diapause, an untypical case of phenotypic plasticity. The hypothesis that Sc. stercoraria, whose generation times are much longer than those of Se. cynipsea, responds to a greater extent to temperature rather than photoperiod only was rejected.

For many insect species, pheromones are important communication tools, but chemical analysis and experimental study can be technically challenging because they require the detection and handling of complex chemicals in small quantities. One drawback of traditional mass spectrometry methods such as gas chromatography mass spectrometry is that whole-body extractions from one to several hundred individuals are required, with the consequence that intra- and interindividual differences cannot be detected. Here, we used the recently introduced UV-LDI MS (ultraviolet laser desorption/ionization mass spectrometry) to profile the ‘osmeterium’ of the sepsid fly Themira superba that is located on the edge of the hind tibia of males. Based on analyses of individual legs, we established that the gland produced a secretion that consisted of oxygenated hydrocarbons and putative isoprenoids. The secretion was first detected 24 h after eclosion, and its transfer to the wings of females during mating was demonstrated using UV-LDI MS. We then tested whether the secretion had an anti-aphrodisiac function, but experimental transfer of the secretion to virgin females did not affect mating success or copulation duration. Throughout the study, UV-LDI MS proved invaluable, because it allowed tracking the natural and experimental transfer of small quantities of pheromones to specific body parts of small flies.

The maintenance of genetic variation in traits under strong sexual selection is a longstanding problem in evolutionary biology. The genic capture model proposes that this problem can be explained by the evolution of condition dependence in exaggerated male traits. We tested the predictions that condition dependence should be more pronounced in male sexual traits and that genetic variance in expression of these traits should increase under stress as among-genotype variation in overall condition is exposed. Genetic variance in female and nonsexual traits should, by contrast, be similar across environments as a result of stabilizing selection on trait expression. The relationship between the degree of sexual dimorphism, condition dependence and additive genetic variance (Va ) was assessed for two morphological traits (body size and relative fore femur width) affecting male mating success in the black scavenger fly Sepsis punctum (Diptera: Sepsidae) and for development time (a nonsexual trait often correlated with body size). We compared trait expression between the sexes for two cross-continental populations that differ in degree of sexual dimorphism (Ottawa and Zurich). Condition dependence was indeed most pronounced in males of the strongly dimorphic Zurich population (males larger), and Va was similar for males and females unless the trait was strongly sex specific and condition dependent. Contrary to prediction, however, Va primarily increased under food limitation in both sexes, and genetic variance in fore femur width was low to nil, perhaps depleted by putatively strong sexual selection. Solely for body size of Zurich males, Va increased more in males than females at limited food, in accordance with the predictions of the genic capture model. Overall therefore, quantitative genetic evidence in support of the model was inconsistent and weak at best.

A standardized bioassay previously developed with ivermectin for the yellow dung fly (Scathophagidae) and the face fly (Muscidae) was applied to test the response of 11 dung fly species to the presumably less toxic parasiticide moxidectin. The results were compared to existing data for the same species tested with ivermectin, albeit two new species (Scathophaga suilla and Musca domestica) were tested here with both the substances. Estimated lethal effect concentrations LC50 at which 50% of the flies died ranged more than tenfold from 0.012 mg moxidectin/kg fresh dung for Sepsis neocynipsea (Sepsidae) to 0.140 mg moxidectin/kg fresh dung for the house fly Musca domestica (Muscidae). In most of the species, we additionally revealed sub-lethal effects at lower moxidectin concentrations in terms of retarded growth and development and reduced body size. Mortality thresholds were about ten times higher for moxidectin than for ivermectin, hence moxidectin is indeed less toxic than ivermectin in absolute terms. Crucially, we obtained strong correlations among the 11 tested fly species in both lethal and sub-lethal responses to the two substances, such that species relatively sensitive to ivermectin were also relatively sensitive to moxidectin. Such correlations are expected if the two substances are structurally related and function in the same manner by disturbing ion channel transport. Methodologically speaking, all species used proved suitable for toxicological testing of parasiticides.

Males of the dung flySepsis cynipsea(L.) (Diptera: Sepsidae) mate guard before copulation by mounting the female's dorsum upon her arrival at the dung. After completing oviposition, females walk off the dung and vigorously shake to attempt to dislodge the guarding male. Only males that succeed in remaining on the female's dorsum will establish genital contact and copulate. Males are smaller than females, and engage wing clamps (specially modified areas on the femur and tibia of each foreleg) to grip the female's wing bases during guarding. The foretibiae of males and females collected at three phases of pairing (prior to struggling, during struggling and in copula) were measured. No differences in the foretibia size of either males or females across the three phases were found. However, the absolute fluctuating asymmetry of the foretibiae of males differed significantly between phases. On average, males paired in copula were the most symmetrical whilst those paired prior to struggling were the least symmetrical. It was concluded that males with more symmetrical foretibiae are indirectly selected as mates by females because of their better competitive ability to grip the female's wing bases and thereby remain on the dorsum of a struggling female during guarding.

Diplogaster coprophila is specifically phoretic on Sepsidae, Sphaeroceridae (Diptera) and also Cercyon species (Coleoptera, Hydrophilidae). Dauerlarvae of D. coprophila were present on 59% of sepsids caught with emergence traps and 13% of those that landed on a cow pat in the field. Both sexes of the flies were infested. In laboratory experiments, using Sepsis punctum as carriers, dauerlarvae were found to aggregate on the fly pupae where they wait motionless until the metamorphosis of the fly is completed. At eclosion, the dauerlarvae move to the abdomen of the fly and enter the genital opening; they are present in the reproductive system of female and male sepsids. Dauerlarvae are transmitted from male to female during copulation. They cannot leave the carriers by their own volition but are placed in the dung by the female fly during oviposition. Phoresy is physiologically obligatory for D. coprophila. Dauerlarvae fail to develop unless they spend some time on a carrier.

The Diptera occurring in various kinds of bird nests were investigated in the Japanese island of Hokkaido from 1985 to 1989. A total of 4623 Diptera, consisting of 23 species of eight families (Anthomyiidae, Fannidae, Muscidae, Calliphoridae, Heleomyzidae, Sphaeroceridae, Sepsidae, and Stratiomyiidae), was reared from 69 natural nests of 13 bird species of eight families (Strigidae, Picidae, Paridae, Sittidae, Sturnidae, Ploceidae, Laniidae, and Columbidae). Of 23 species of flies reared, 12 were new records from bird nests. An ectoparasite of birds,Protocalliphora maruyamensisShinonaga and Kano, frequently emerged from the nests of hole-nesting birds of Paridae, Sturnidae, and Ploceidae, but it did not emerge from the nests of the hole-nesting birdsPicoides majorandP.minorof Picidae. The nests ofStreptopelia orientalisyielded the most species of flies (12).Myospila meditabunda(Fabricius) andMydaea urbana(Meigen), whose larvae are predacious, proved able to live in the bird nests. Among the scavengers,Potamia littoralis(R.-D.) was the most dominant and occurred in the nests of nine of 10 hole-nesting bird species. Sphaerocerid, sepsid, and stratiomid flies occurred in the nests ofS.orientalisin which large quantities of bird feces had accumulated. The factors concerning the occurrence of flies in the nests are discussed.

The result of a phylogenetic analysis of the Sepsidae based on larval characters is presented. It is shown that cyclorrhaphan larvae can be as rich a source of characters as Nematocera immatures when investigated using an SEM. The cladistic analysis comprised fifty-two species in sixteen genera of the Sepsidae and five outgroup species and used fifty-seven morphological characters. It found seven parsimonious trees which only differed with respect to the arrangement of some species within the genus Themira. The basal dichotomies of the phylogenetic trees are particularly well supported, indicating the conservative nature of larval characters. Orygma is confirmed as the sister group of all the remaining sepsids, the Sepsinae. There is good larval evidence that Ortalischema is the sister group of all remaining Sepsinae and that the Toxopodinae constitute an early radiation within the Sepsidae. According to larval data, some genera are paraphyletic (Themira, Palaeosepsis), but adult characters appear to contradict these findings. Among the traditionally recognized higher taxa within the Sepsidae, Hennig's Themira species-group and Steysbal's Sepsini have to be rejected as polyphyletic. However, Hennig's Sepsis species-group is confirmed as monophyletic and will probably constitute one major element of a future phylogenetic system of the Sepsidae. States of the strongly modified fore-legs of some adult sepsid males are mapped onto the phylogenetic tree, largely confirming Sulc's ideas about the evolution of these features. The origin and evolution of male sternites with brushes and a gland on the tibiae of the males (‘osmeterium’) are discussed. Whereas adult characters point to a sister-group relationship between the Sepsidae and the Ropalomeridae, larval characters appear to indicate a sister-group relationship between the Coelopidae and the Sepsidae. The evidence for both hypotheses is critically evaluated.

Sexual size dimorphism (SSD) varies widely across and within species. The differential equilibrium model of SSD explains dimorphism as the evolutionary outcome of consistent differences in natural and sexual selection between the sexes. Here, we comprehensively examine a unique cross-continental reversal in SSD in the dung fly, Sepsis punctum. Using common garden laboratory experiments, we establish that SSD is male-biased in Europe and female-biased in North America. When estimating sexual (pairing success) and fecundity selection (clutch size of female partner) on males under three operational sex ratios (OSRs), we find that the intensity of sexual selection is significantly stronger in European versus North American populations, increasing with male body size and OSR in the former only. Fecundity selection on female body size also increases strongly with egg number and weakly with egg volume, however, equally on both continents. Finally, viability selection on body size in terms of intrinsic (physiological) adult life span in the laboratory is overall nil and does not vary significantly across all seven populations. Although it is impossible to prove causality, our results confirm the differential equilibrium model of SSD in that differences in sexual selection intensity account for the reversal in SSD in European versus North American populations, presumably mediating the ongoing speciation process in S. punctum.

There is accumulating evidence that criminals wrap dead bodies in an attempt to conceal evidence. To anticipate the forensic implications of this phenomenon, we examined whether flies that are naturally associated with cadavers exhibit a delay in attendance or differ in species composition and abundance patterns because of the presence of wrapping material. Wrapped and exposed carcasses of dead monkeys placed in an oil plantation in Kedah, Malaysia, were visited over 50 d. On daily visits to each of the six carcasses, visiting adult flies were sampled using hand nets. Flies of 12 families were encountered. Calliphoridae (Chrysomya rufifacies Macquart and C. megacephala (F.) was the most prevalent family, followed by Sphaeroceridae. Some families tended to be more abundant in WRCs (i.e., Calliphoridae, Muscidae, and Phoridae), whereas others (i.e., Piophilidae, Sepsidae, and Psychodidae) were more prevalent in exposed carcasses. Wrapping delayed the arrival of all fly species encountered, with delays varying from 1 to 13 d depending on species. Wrapping did not affect species composition of flies, but prolong the occurrence of some species. The results of the current study emphasize the need to take into consideration the presence of a wrap when estimating postmortem interval.