taxonID	type	description	language	source
814387FFFFB5FFFD048B3E9F5665F8C9.taxon	description	Subgenus Argoravinia (s. s.): male with 5 – 6 fronto-orbital setulae; epandrium with a lateral apophysis *; vesica superficially bifid; female T 6 entire; female epiproct with one seta. Subgenus Raviniopsis Townsend: male with 7 – 12 fronto-orbital setulae; epandrium without a lateral apophysis; vesica deeply bifid; female T 6 divided; female epiproct with two setae.	en	Buenaventura, Eliana, Pape, Thomas (2018): Phylogeny, evolution and male terminalia functionality of Sarcophaginae (Diptera: Sarcophagidae). Zoological Journal of the Linnean Society 183 (4): 808-906, DOI: 10.1093/zoolinnean/zlx070, URL: https://academic.oup.com/zoolinnean/article/183/4/808/4757488
814387FFFFB3FFFA071338365585FE78.taxon	description	Subgenus Cistudinomyia: posterior postgenal setulae white; epandrium reddish, usually the same colour as syntergosternite 7 + 8; distiphallus without paraphallic distal expansions. Subgenus Dexosarcophaga: genal and postgenal setulae generally black; white setulae, when present, are very scarce and restricted to the posteriormost part of the postgena; epandrium blackish, usually the same colour as syntergosternite 7 + 8 *; distiphallus with paraphallic distal expansions.	en	Buenaventura, Eliana, Pape, Thomas (2018): Phylogeny, evolution and male terminalia functionality of Sarcophaginae (Diptera: Sarcophagidae). Zoological Journal of the Linnean Society 183 (4): 808-906, DOI: 10.1093/zoolinnean/zlx070, URL: https://academic.oup.com/zoolinnean/article/183/4/808/4757488
814387FFFFB0FFF804B9392C5035FEA2.taxon	description	Subgenus Dodgeisca: male mid-femur with a ctenidium of rounded spines (circular cross section); pregonite straight, sclerotized, as long as phallus; hillae tube-like distally *.	en	Buenaventura, Eliana, Pape, Thomas (2018): Phylogeny, evolution and male terminalia functionality of Sarcophaginae (Diptera: Sarcophagidae). Zoological Journal of the Linnean Society 183 (4): 808-906, DOI: 10.1093/zoolinnean/zlx070, URL: https://academic.oup.com/zoolinnean/article/183/4/808/4757488
814387FFFF8DFFC507133D83501FFBB9.taxon	description	Subgenus Pacatuba: male mid-femur with a ctenidium of rounded spines (circular cross section); wing vein R 1 bare dorsally; metasternum setulose. Subgenus Sarcofahrtiopsis: male mid-femur without a ctenidium; wing vein R 1 setulose dorsally; metasternum with reduced setosity *.	en	Buenaventura, Eliana, Pape, Thomas (2018): Phylogeny, evolution and male terminalia functionality of Sarcophaginae (Diptera: Sarcophagidae). Zoological Journal of the Linnean Society 183 (4): 808-906, DOI: 10.1093/zoolinnean/zlx070, URL: https://academic.oup.com/zoolinnean/article/183/4/808/4757488
814387FFFF8AFFC204B93CC855F9FB76.taxon	description	Subgenus Carinoclypeus: facial carina parallel in full length to frontogenal suture *; cercal prong without a proximal tuft of long black setae. Subgenus Udamopyga (s. s.): facial ridge with dense setosity on lower 0.50; cercal prong with a proximal tuft of long black setae.	en	Buenaventura, Eliana, Pape, Thomas (2018): Phylogeny, evolution and male terminalia functionality of Sarcophaginae (Diptera: Sarcophagidae). Zoological Journal of the Linnean Society 183 (4): 808-906, DOI: 10.1093/zoolinnean/zlx070, URL: https://academic.oup.com/zoolinnean/article/183/4/808/4757488
814387FFFF86FFCD04B93E1B50A2FBAD.taxon	description	It is noteworthy that the four genera of theTricharaea grade share a fair number of features not found outside this group, yet they emerge as paraphyletic in our analysis. The following shared character states would appear particularly relevant in this context: proclinate fronto-orbital setae in males (‘ pc’ in Fig. 41), notopleuron without subprimary setae (‘ nt’ in Fig. 41 B – D), two katepisternal setae, postalar wall bare, wing vein R 4 + 5 with dorsal setulosity reaching crossvein r-m, ST 5 with posterior margin straight or with a shallow concavity (Fig. 42 A, B), ST 5 with a central patch of setae (Fig. 42 A), vesica divided into a proximal and a distal section (Fig. 25 A – D), vesical arm-shaped lever elongated to very elongated ventrally (Fig. 25 A – D), vesical arm-shaped lever with a hammer-shaped or bilobed to oval apex (Figs 19 H, 27 A, 28 H, J) and distal section of the vesica globose, with small denticles (Figs 19 E, 28 G, H, 39 H). The last five character states are found only in species of the Tricharaea grade. The genera Bahamiola and Sarcofahrtiopsis (including Pacatuba) do not form a monophyletic group, but they share a vesical arm-shaped lever very elongated ventrally (Figs 15 H, 19 E, 28 H) and a hood-shaped juxta with a denticulated lateral margin that is enlarged ventrally to form a capsule-like structure (Figs 15 G, 19 E – G, 28 H, J, 30 A, B). Additional characters and a larger sample of outgroup taxa will be a proper test of this topology, and therefore of the polarity of the character transformation series involved in the evolution of the ‘ lower’ sarcophagines. The monophyly of Tricharaea was previously supported by molecular data (Piwczyński et al., 2014), but here it is also supported by four autapomorphies: epandrium brownish (not reddish), vesical arm-shaped lever elongated (Figs 25 A, 27 A), juxta smooth laterally and wrinkled medially (Figs 27 A, 39 H), juxta funnelshaped (Fig. 39 H). Within the Sarcophaginae, two plesiomorphic character states are shared by the three taxa of Tricharaea and Paramacronychiinae: postgena angled in lateral view (Fig. 41 D, E), and sparse, weak anepimeral setulae (‘ as’ in Fig. 41 D). In the handmade cladogram of the tribe Sarothromyiini, Lopes (1990) argued for the monophyly of Tricharaea based on its species sharing spherical spermathecae. Later, Pape (1996) used this character state plus five features of male terminalia structures, three of female terminalia and one of the puparium, to diagnose the genus Tricharaea. Pape’s (1996) male character states were: (1) male with at least one strong proclinate orbital seta, (2) postalar wall bare, (3) metasternum setulose, (4) male ST 5 with a central patch of setae, (5) terminalia brownish (not red), (6) spermathecae spherical, (7) female with an epiproct and (8) puparial spiracles not in a recession. Except for female and larval character states 6 – 8, all others were included here, and only character states 1 and 5 (slightly modified) were found to be autapomorphic for this genus. However, all of Pape’s (1996) character states and the two plesiomorphic and one autapomorphy found in the present study are used to diagnose this genus. With a single species, the genus Sarcofahrtiopsis was described by Hall (1933) based on ST 5 not having a cleft. Dodge (1965 b) added more character states to the diagnosis of this genus, such as the hind coxa bare posteriorly, wing vein R 1 setulose and proclinate orbital setae present in males. Later, Lopes (1990) suggested the setulose wing vein R 1 and the long and bristly pregonite as synapomorphies; however, the first character state is also shared with genera such as Helicobia, Malacophagomyia (including Dodgeisca), Nephochaetopteryx, Panava, Promayoa, Rafaelia, among others, and the second character state does not diagnose Sarcofahrtiopsis, as it is not present in all species of the genus. Pape’s (1996) diagnosis included the mentioned character states plus the following: notopleuron with subprimary setae, postalar wall bare, metasternum bare, third costal sector of wing bare ventrally, male ST 5 with a central patch of setae, terminalia usually black, spermathecae elliptical and female without an epiproct. Finally, Mello-Patiu & Pape (2000) discussed all these features and suggested a list of 16 character states as a generic diagnosis of Sarcofahrtiopsis, highlighting the reduced metasternal setosity and the slender and elongated parameral (= postgonal) apodeme as autapomorphies. From these, the slender parameral apodeme should probably be removed as an autapomorphy, since this structure is not elongated in Sarcofahrtiopsis thyropteronthos Pape, Dechmann & Vonhof, 2002 (Pape, Dechmann & Vonhof, 2002). Here, the 13 male character states of Mello-Patiu & Pape (2000) were analysed and only the reduction in the setosity of the metasternal area came out as autapomorphic for Sarcofahrtiopsis. Neither the monospecific genus Pacatuba nor the polyspecific genus Bahamiola of the classification of Pape (1996), here represented by a single species only, were found to possess any autapomorphies. However, Pacatuba and Sarcofahrtiopsis share one autapomorphy, vesical arm-shaped lever very elongated (twice its full length). The clade of Sarcofahrtiopsis (including Pacatuba) received aBS and weak JK support; however, Pacatuba shares 10 out of the 13 male character states listed by Mello-Patiu & Pape (2000) to define Sarcofahrtiopsis. Therefore, we suggest Pacatuba as a new synonym of Sarcofahrtiopsis. Consequently, we present a new generic diagnosis for Sarcofahrtiopsis, which is divided into the two subgenera Pacatuba, new status, and Sarcofahrtiopsis (s. s.), for which we also include subgeneric diagnoses.	en	Buenaventura, Eliana, Pape, Thomas (2018): Phylogeny, evolution and male terminalia functionality of Sarcophaginae (Diptera: Sarcophagidae). Zoological Journal of the Linnean Society 183 (4): 808-906, DOI: 10.1093/zoolinnean/zlx070, URL: https://academic.oup.com/zoolinnean/article/183/4/808/4757488
814387FFFF85FFCA07133909557FFC9B.taxon	description	Species of Nephochaetopteryx are here included for the first time in a phylogenetic study. The phylogenetic affinity between this genus and Tricharaea, and also the position of these two genera within the ‘ lower’ Sarcophaginae, was suggested by Lopes (1983) on the basis of these genera sharing first-instar larval character states such as a vestigial labrum [= mandible in Lopes (1983)] and the dorsal bridge [= clypeal arch in Lopes (1983)] situated posterior to the parastomal bar [= paraclypeal phragma in Lopes (1983)]. Interestingly, in the same study, Lopes also considered the clade composed of Oxysarcodexia and Ravinia as sister group of Nephochaetopteryx due to these genera also sharing the first-instar larval character states mentioned above. Later, in his phylogenetic study of the Sarcophaginae males with proclinate orbital setae, Lopes (1990) placed Nephochaetopteryx as sister to Rettenmeyerina due to these two genera sharing the distiphallus articulated with the basiphallus. However, here we found that only Rettenmeyerina possesses a fully developed hinge between basi- and distiphallus (Fig. 28 D), while Nephochaetopteryx generally has a desclerotized strip or a superficial hinge and only dorsally (Fig. 29 H). Nephochaetopteryx was placed together with Bahamiola, Sarcofahrtiopsis, Rettenmeyerina and Tricharaea in the tribe Sarothromyiini by Lopes (1969 a), due to all males of these genera having proclinate fronto-orbital setae. Males with proclinate fronto-orbital setae are largely confined to genera within the ‘ lower’ Sarcophaginae, with few exceptions like in Duckemyia, two species of Lepidodexia, species of Panava, one species of Tripanurga and a few species of Helicobia. The sister-group relationship between Oxysarcodexia and Ravinia was highlighted already by Roback (1954), who pointed out similarities in phallic structures such as the lack of a juxta (i. e. the lack of juxtal hinge) and the presence of the acrophallic levers [= dorsal rods in Roback (1954)]. This was later followed by Downes (1955), who added larval and female traits in support of this relationship. Using Buenaventura & Pape’s (2015) broader definition of the juxta, all genera of Sarcophaginae possess this structure. The hypothesis of a sister-group relationship between Oxysarcodexia and Ravinia was corroborated by Pape (1994) and Giroux et al. (2010) based on morphological data. One molecular-based phylogenetic analysis found strong support for this relationship (Stamper et al., 2012), while two others (Kutty et al., 2010; Piwczyński et al., 2014) did not, although the last two analyses showed low branch supports. In our analysis, the clade (Oxysarcodexia + Ravinia) is supported by the homoplasious character state of a ctenidium of flattened spines (also found in Mecynocorpus and most Paramacronychiinae). This clade is also supported by the first-instar larval character state of festoon-like oral ridges (Downes, 1955; Lopes, 1983; Leite & Lopes, 1987; Lopes & Leite, 1987; Pape, 1996). Forthefirsttime, themonophylyof Nephochaetopteryx is tested in a modern phylogenetic context, and its monophyly received strong JK support and has three male external autapomorphies: apical part of wing membrane between veins R 2 + 3 and C fumose, mid-tibia without antero-dorsal setae and hind coxa with strong posterior setae. None of these character states were used in the original description of Nephochaetopteryx by Townsend (1934), but later Dodge (1968 a) provided a first diagnosis for this genus, where he included the following character states: mid-tibia with neither antero-dorsal nor antero-ventral setae, wing vein R 1 setulose and arista plumose on basal three-fifths. Later, Lopes (1990), in his handmade cladogram, included the second character state of Dodge (1968 a) plus the reduction of the female eighth tergite in his ‘ list of synapomorphies’ of Nephochaetopteryx. Lastly, Pape (1996) provided a diagnosis including 14 character states, 13 of which were analysed here, and three of which were found to be autapomorphies for this genus. These three character states, in combination with some of Pape’s (1996) other character states, are used here to diagnose Nephochaetopteryx. The monophyly of Oxysarcodexia was already inferred in non-cladistic studies (Lopes, 1943, 1983; Roback, 1954), and later confirmed by phylogenetic analyses using both morphological (Giroux et al., 2010) and molecular (Stamper et al., 2012; Piwczyński et al., 2014) characters. In our analysis, Oxysarcodexia is supported by two autapomorphies: (1) paraphallus antero-proximally with a paraphallic triangular expansion proximal to the vesica (‘ pte’ in Fig. 18 H) and (2) juxta with a proximal convex membranous expansion (‘ jce’ in Fig. 18 H). The first character state was recognized as diagnostic for this genus in previous studies (Lopes, 1946; Dodge, 1966; Giroux et al., 2010). The second character state was first described by Lopes (1946) in his detailed revision of Oxysarcodexia, where the species descriptions used mostly male terminalia characters, such as the vesica, since this structure has a remarkable morphological diversity in this genus. In a subsequent work, Lopes (1975 b) erected the subtribe Oxysarcodexiina, which he defined with a reduced list of diagnostic character states when compared to his earlier work. A selection of eight of Lopes’s (1946, 1975 b) character states was listed in a more recent diagnosis for this genus (Pape, 1996), which, however, did not include the character states found as autapomorphic here. Subsequent authors used these two character states in descriptions of new species (Soares & Mello-Patiu, 2010) and in morphological comparative studies (Silva & Mello-Patiu, 2008). Besides the two autapomorphies and some homoplasies found in our analysis, Oxysarcodexia is here diagnosed with three additional external male character states and two first-instar larval character states as suggested by previous studies. The monophyly of Ravinia was suggested by Roback (1954) and Lopes (1983), and recently both morphology-based (Giroux et al., 2010) and molecular-based (Stamper et al., 2012; Piwczyński et al., 2014) phylogenetic studies have corroborated this hypothesis. Here, five autapomorphies supported the monophyly of Ravinia: juxta hood-shaped, partially wrinkled and slightly swollen (Figs 10 C, D, 14 G, H), hillae distally blunt (Fig. 10 C) or pointed (Fig. 14 E), vesica narrow and flake-shaped (Figs 10 C, D, 14 G), vesical arm-shaped lever straight proximally (Fig. 35 D) and distal section of the vesica flattened or reduced (Fig. 10 D). Giroux et al. (2010) found the presence of hillae as the only autapomorphy for this genus, but in our definition this structure is also found in an additional 15 genera. However, the hillae in Ravinia are highly specialized in comparison to those found in other genera. Specifically, hillae with a membranous bladder (Figs 10 C, D, 14 G, H) and a groove (Fig. 10 B), as described by Giroux et al. (2010), are only found in some species of this genus. The importance of the hillae in the definition of Ravinia was already mentioned by Roback (1954), who also inferred the origin of the acrophallic levers [= acrophallic bars in Roback (1954)] in other taxa [acrophallic levers originated in the ancestor of all Sarcophaginae (clade 4 in Fig. 2 A) according to our analysis] before the emergence of the Ravinia lineage. Five autapomorphies supporting the monophyly of Ravinia are used to diagnose this genus in combination with other male structures and two larval character states.	en	Buenaventura, Eliana, Pape, Thomas (2018): Phylogeny, evolution and male terminalia functionality of Sarcophaginae (Diptera: Sarcophagidae). Zoological Journal of the Linnean Society 183 (4): 808-906, DOI: 10.1093/zoolinnean/zlx070, URL: https://academic.oup.com/zoolinnean/article/183/4/808/4757488
814387FFFF82FFC904B93E195697FCE5.taxon	description	Roback (1954) considered Cistudinomyia as part of the subtribe Raviniina, Dodge (1968 b) considered Dexosarcophaga as closely related to Oxysarcodexia, while Lopes (1969 a, 1975 b, 1983) did not include Cistudinomyia in his tribal array of the Sarcophaginae, but he placed Dexosarcophaga in the tribe Cuculomyiina, Oxyvinia in Raviniini and Rettenmeyerina in Sarothromyiini. Giroux et al. (2010) included Cistudinomyia, Dexosarcophaga and Oxyvinia in their taxon sample and found a weakly supported clade (( Dexosarcophaga + Oxyvinia) + (Cistudinomyia + other Sarcophaginae )) using morphological characters. The molecular studies of Kutty et al. (2010) and Piwczyński et al. (2014) included Dexosarcophaga and recovered the topologies (Dexosarcophaga + (Argoravinia + Blaesoxipha )) and (Dexosarcophaga + Argoravinia), respectively, both with low branch support. Lopes (1969 a) placed Rettenmeyerina together with Bahamiola, Sarcofahrtiopsis and Tricharaea in the tribe Sarothromyiini on the basis of these genera sharing proclinate fronto-orbital setae in the male. Here, Rettenmeyerina is diagnosed only by homoplasies, as we found no autapomorphies for this genus. The presence of a desclerotized area between the paraphallus and the juxta in Rettenmeyerina is relevant for defining this genus. Rettenmeyerina emerges as sister taxon to the remaining ‘ higher’ Sarcophaginae, which has (Oxyvinia + Dexosarcophaga [including Cistudinomyia]) as sister clade of the remaining Sarcophaginae species (Fig. 2 A). The presence of proclinate fronto-orbital setae in the male is a plesiomorphic feature in the Tricharaea grade, which means that the absence of male proclinate fronto-orbital setae in the ancestor of the ‘ higher’ sarcophagines (excl. of Rettenmeyerina) has to be considered an apomorphic reversal. Male proclinate fronto-orbital setae, i. e. male and female with the same frontal chaetotaxy, are of very sporadic occurrence in the Calyptratae, and there is to our knowledge no other instance where the presence of male proclinate orbital setae has been hypothesized as a reversal.	en	Buenaventura, Eliana, Pape, Thomas (2018): Phylogeny, evolution and male terminalia functionality of Sarcophaginae (Diptera: Sarcophagidae). Zoological Journal of the Linnean Society 183 (4): 808-906, DOI: 10.1093/zoolinnean/zlx070, URL: https://academic.oup.com/zoolinnean/article/183/4/808/4757488
814387FFFF81FFDC048B3BB054BEF8EA.taxon	description	The clade (Malacophagula + Rafaelia) is supported by four homoplasies: parafacial plate with strong setae (Fig. 42 F), male hind tibia with apical postero-ventral setae well differentiated, a median stylus moderately elongated and a demarcated juxta with a hinge or a desclerotized strip between the juxta and the remaining distiphallus (Figs 14 B, 17 G). Species of the genera Malacophagula and Rafaelia have never been studied with modern phylogenetic methods, but the tribal classification based on first-instar larval character states proposed by Lopes (1983) included these genera together with species of Lepidodexia and Titanogrypa in the tribe Johnsoniini. Mello-Patiu & Azevedo (1998) also highlighted similarities observed by Lopes (1983) in the median and lateral styli of genera Malacophagula and Rafaelia and differences in head morphology for which we found support here. The vesica in these genera requires deeper study, as it could carry informative characters for defining the two genera and reconstructing their species-level phylogenetic relationships. The monophyly of Malacophagula is strongly supported by five autapomorphies: head rounded in profile (Fig. 42 F), first flagellomere shortened (Fig. 42 F), lunule widened, postgena swollen (Fig. 42 F) and lower calypter rounded (Fig. 43 A). One autapomorphy and three homoplasies supported the monophyly of Rafaelia, which received moderate branch support (Fig. 2 B). The only autapomorphy for this genus was hypophallus weakly sclerotized, with only the very apex of the vesica sclerotized. Species of Rafaelia have a hypophallus that is mostly membranous, globose and well developed, while the paraphallus consists of a thin, sclerotized dorsal plate (Figs 20 F, 40 C, D), which is a rare condition in Sarcophaginae. Roback (1954) considered Argoravinia as part of the Johnsonia Coquillett group, which included species of Lepidodexia, Emblemasoma and Helicobia, although he explicitly affirmed this as a tentative placement since he did not find any resemblance of the phallic structures of this genus to those of any other Sarcophaginae. In his classification based on first-instar larval character states, Lopes (1983) included Argoravinia in the Sarcodexiina group together with species of Peckia, Helicobia and Lipoptilocnema. Molecular studies including only few Neotropical genera have marginally touched upon the phylogenetic position of Argoravinia with regard to other Sarcophaginae (Kutty et al., 2010; Piwczyński et al., 2014). These studies showed conflicting relationships for this genus, either as sister to Blaesoxipha setosa (Salem, 1938) with moderate to strong support (Kutty et al., 2010), or to Dexosarcophaga transita Townsend, 1917 with no branch support (Piwczyński et al., 2014). In our analysis, which includes a larger taxon sample than previous phylogenetic studies on Sarcophaginae, Argoravinia emerges as sister to Malacophagomyia (including Dodgeisca) due to these taxa sharing three autapomorphies: (1) head profile with squared anterior and posterior genal corners, (2) paraphallic lateral expansions (Figs 15 A – C, 16 C, 17 A) and (3) median stylus greatly elongated (Figs 15 B, E, 16 D, 17 A, B, D). The delimitation and monophyly of Argoravinia was revised by Pape (1990) but is here explicitly tested for the first time, and it received aBS and strong JK support. This genus is supported by six autapomorphies: (1) stem of wing vein R 2 + 3 + 4 + 5 with ventral setulae elongated, (2) pregonite proximally narrow and distally wide, (3) hillae convoluted (Fig. 15 B – F), (4) capitis as a smooth, rounded lobe, proximally swollen, (5) median stylus S-shaped (Fig. 15 B, E) and (6) juxta very small to vestigial (Fig. 15 E). Some of these character states were previously included in the generic diagnoses for Argoravinia (Lopes, 1976 a; Pape, 1990, 1996; Carvalho-Filho & Esposito, 2012). For example, Lopes (1976 a) mentioned the long styli with a conspicuous free base, and a ‘ median process of glans’ with a long slender ‘ apophysis’, which partially correspond to our character states of the hillae and capitis, respectively. Similarly, in the diagnosis of Argoravinia, Pape (1990, 1996) included stem of wing vein R 2 + 3 + 4 + 5 with ventral setulae elongated, and the median stylus S-shaped, both found here as autapomorphic for this genus. More recently, Carvalho-Filho & Esposito (2012) diagnosed this genus based on nine character states, but only the vestigial juxta emerged as autapomorphic, and all others as homoplastic in the present analysis. Due to their utility for sorting Argoravinia species from other genera, most of the character states proposed by the above-mentioned authors are included in our diagnosis. Finally, the monophyly of the subgenera proposed by Carvalho-Filho & Esposito (2012) is partially supported by our phylogeny, as we recovered a monophyletic Argoravinia (s. s.), but as only a single species of Raviniopsis was included, its possible monophyly remains untested (Fig. 2 A). The subgeneric classification of the genus Argoravinia proposed by Carvalho-Filho & Esposito (2012) was supported by the following character states: (1) setulae colour on the gena as black for Argoravinia (s. s.) and white for Raviniopsis, but here scored as gena and postgena having at least some setulae white for all Argoravinia species; (2) number of fronto-orbital setulae, which was not included here; (3) bending of the cerci and presence / absence of a cluster of spines apically, which we considered as two separate characters and scored cerci as straight or almost straight for all Argoravinia species since the ‘ bent’ condition is only observed in taxa of the Blaesoxipha clade, and the cercal spines as ‘ a cluster’ were not included here; (4) male epandrium with a lateral apophysis for Argoravinia (s. s.) or without for Raviniopsis, which was included and supported the monophyly of Argoravinia (s. s.) in our phylogenetic analysis; (5) vesica bifid for Argoravinia (s. s.) or composed of two separated lobes for Raviniopsis, which is here scored as bifid for all Argoravinia species, since species that appear to have two separated vesical lobes, might actually have the lobes fused at the base; (6) shape of the female T 6, which was not included here; and (7) female with one seta on the epiproct in Argoravinia (s. s.) or two seate in Raviniopsis, which was not included here. Thus, our results support the subgeneric classification by Carvalho-Filho & Esposito (2012), since the presence of an epandrial lateral apophysis in species of Argoravinia (s. s.) was found as autapomorphic for this subgenus. Species of Malacophagomyia are here included for the first time in a phylogenetic study. Lopes (1969 a, 1983) implied a phylogenetic affinity of this genus to genera such as Titanogrypa, Panava, Dexosarcophaga and Udamopyga, but this is not supported by our results. The three studies providing a diagnosis for this genus (Lopes, 1966; Pape, 1996; Mulieri & Mello-Patiu, 2013) highlighted the remarkably elongated median stylus and the conspicuous juxta, which are characteristic for all species of Malacophagomyia. In at least two (i. e. Pape, 1996; Mulieri & Mello-Patiu, 2013) of these studies, the authors agree on the following consensus list of diagnostic character states: (1) postalar wall setulose, (2) male mid-femur without a ctenidium, (3) wing vein R 1 setulose dorsally, (4) third costal sector of wing setulose ventrally, (5) pregonite with membranous area along the ventral margin and near the bent apical part, (6) acrophallus with median stylus greatly elongated and curved (Fig. 17 A, B, D) and (7) juxta arching over the lateral styli (Fig. 17 A, B, D). Interestingly, the most remarkable character states (6 and 7) are shared with the species Dodgeisca paramerata Rohdendorf, 1971 (Fig. 16 C, D), the only known species of Dodgeisca, which also shares with Malacophagomyia character states 1, 3, 4 of this consensus list. In addition, according to the most recent revision of Malacophagomyia (Mulieri & Mello-Patiu, 2013), not all species of this genus possess character state 5, which leaves only character state 2 (male mid-femur without a ctenidium) as a difference between Dodgeisca and Malacophagomyia. Besides that, in their revision of the latter genus, Mulieri & Mello-Patiu (2013) highlighted the cerci fused along their entire length and the spine-like setae on ST 4 as possible autapomorphies of Malacophagomyia. Both of these character states are also present in D. paramerata. Mulieri & Mello-Patiu (2013) also included the absence of a vesica, the presence of harpes and arms of the lateral styli as part of their diagnosis of Malacophagomyia. According to our observations, both Malacophagomyia and Dodgeisca possess a broad and flat vesica (Figs 16 C, D, 17 A, B, D), which, however, is not as prominent as in other sarcophagines. Also, the ‘ arms of the lateral styli’ described by Mulieri & Mello-Patiu (2013) are consistent with our definition of hillae, while the structures considered as harpes by these authors do not follow our definition for that structure. Consequently, the ‘ arms of the lateral styli’ (Mulieri & Mello-Patiu, 2013) are homologized with the hillae (Figs 16 C – E, 17 A, B), and their ‘ harpes’ with the paraphallic lateral expansions (Figs 16 C, 17 A). In addition to the synapomorphies mentioned above, we found Malacophagomyia and Dodgeisca to share the presence of two pointed processes on the juxtal apex (Figs 16 D, 17 B), and distal part of hillae membranous. Based on all the above, we suggest Dodgeisca as a new junior synonym of Malacophagomyia, and we maintain it and give it a new status as a subgenus of the latter genus.	en	Buenaventura, Eliana, Pape, Thomas (2018): Phylogeny, evolution and male terminalia functionality of Sarcophaginae (Diptera: Sarcophagidae). Zoological Journal of the Linnean Society 183 (4): 808-906, DOI: 10.1093/zoolinnean/zlx070, URL: https://academic.oup.com/zoolinnean/article/183/4/808/4757488
814387FFFF94FFDA07013E3F5480F833.taxon	description	The genera of the Blaesoxipha clade share four apomorphic character states: (1) male with abdominal ST 5 cleft with subparallel sides (Fig. 43 B, C), (2) distiphallus not surrounding the acrophallus, styli entirely exposed (except in Comasarcophaga and Spirobolomyia) (Figs 12 A, 23 F, 28 E, 29 D, 30 D, 34 D, 36 G), (3) juxta partially to entirely fused to acrophallic structures (Figs 12 B, 22 E, 30 C, 35 H, 39 E) and (4) juxta straight (Figs 35 H, 39 E). Three additional autapomorphies that evolved in the ancestor of this clade, but which have subsequently become reduced or modified in some of these genera, are: distal margin of juxta with spine-like processes (Figs 30 D, 34 D – F), which evolved into distal margin smooth in clades 42 and 51 (Figs 11 H, 28 E, 35 H, 39 E); lateral styli collapsed with no outlet (Figs 23 J, 28 F, 29 B, 30 E, 34 E, 36 H, 39 F), which is reversed in clades 43 and 51 where a sperm outlet is found (Figs 11 H, 20 D, 21 I, 39 B – D); and lateral styli plate-like, with digitate margins or finger-shaped processes (Figs 23 F, 29 E, 30 C, 34 E, 36 G), which are reversed in clades 45, 48 and 51 where the lateral styli are tube-shaped (Figs 11 H, 12 B, 21 I, 22 E, 39 C). Some branches within the Blaesoxipha clade had low supports, and alternative topologies were retrieved differing in the position of the paraphyletic Table 1.	en	Buenaventura, Eliana, Pape, Thomas (2018): Phylogeny, evolution and male terminalia functionality of Sarcophaginae (Diptera: Sarcophagidae). Zoological Journal of the Linnean Society 183 (4): 808-906, DOI: 10.1093/zoolinnean/zlx070, URL: https://academic.oup.com/zoolinnean/article/183/4/808/4757488
814387FFFFFCFFB307013CA356AAFC63.taxon	description	Two autapomorphies support Tulaeopoda: the posterior surface of the male hind trochanter with a postero-median pad of short setae (position as no. 6 in Fig. 45) and male abdominal ST 3 with two patches of dense, erect, black setae. Contrary to what was suggested by Pape (1996), species of Tulaeopoda possess well-developed, tubular lateral styli (Fig. 27 I, J).	en	Buenaventura, Eliana, Pape, Thomas (2018): Phylogeny, evolution and male terminalia functionality of Sarcophaginae (Diptera: Sarcophagidae). Zoological Journal of the Linnean Society 183 (4): 808-906, DOI: 10.1093/zoolinnean/zlx070, URL: https://academic.oup.com/zoolinnean/article/183/4/808/4757488
814387FFFFFBFFB2048B39515542FE6A.taxon	description	Roback (1954) placed Metoposarcophaga Townsend (= Tripanurga) and genera such as Rafaelia and Boettcheria in the subtribe Boettcheriina. Lopes (1969 a) placed Carinoclypeus, Tripanurga and Udamopyga in the tribe Sarcophagini, but in a subsequent study (Lopes, 1983) he included Tripanurga in Sarcophagini, and Udamopyga in Cuculomyiini. None of these proposals had been consistently tested, as no study had included representative species of these genera. Stamper et al. (2012) found Tripanurga importuna (Walker, 1849) to be sister to the genus Boettcheria, which somehow supports Roback (1954) in placing Tripanurga and Boettcheria in the subtribe Boettcheriina. The sister-group relationship between Tripanurga and Boettcheria received high branch support in Stamper et al. ’ s (2012) phylogeny. Our taxon sample is much more extensive than that analysed by Stamper et al. (2012), as we included multiple species of Boettcheria and Tripanurga. However, the low support for Tripanurga as sister to Udamopyga (including Carinoclypeus) leaves this sister-group relationship as tentative. Future analyses are needed to test which of these alternative topologies is best corroborated. Pape (1990) proposed a broad concept of the genus Tripanurga by including Erucophaga Reinhard, Metoposarcophaga, Zygastropyga Townsend and other genera as synonyms. Pape (1990, 1996) diagnosed Tripanurga with seven character states: (1) male cercus with prong bent backwards, (2) ejaculatory apodeme large, (3) parameral (= postgonal) seta slightly flattened, (4) phallus with an epiphallus-like process at base, (5) basiphallus elongated and narrow, (6) distiphallus compact and globular and (7) ventral margin of distiphallus with fringe of filiform processes. Character state 2 is not included here due to difficulties of coding other taxa; 1, 5 – 7 are reinterpreted, and 3 and 4 came out as autapomorphies. In our phylogenetic analysis Tripanurga is monophyletic, supported by five autapomorphies: (1) male abdominal T 5 with ventral margin pointed (arrows in Fig. 40 E), (2) epandrium higher than wide in lateral view (‘ epd’ in Fig. 40 F), (3) postgonal seta slightly compressed (arrows in Fig. 27 F, G), (4) basiphallus proximally with a dorsal epiphallus-like process (‘ ep’ in Fig. 27 B), (5) vesica with vesical lateral arms (‘ vla’ in Fig. 27 B – E), each with an inner denticulated process (‘ vdp’ in Fig. 27 E). The genus Udamopyga (including Carinoclypeus) is supported by three autapomorphies: (1) posterior margin of the male abdominal ST 5 with a slight undulation halfway between the angle and the tip of the V, and a rounded distal expansion (Fig. 40 G), (2) cercal prongs fused at least halfway to tip and (3) vesica composed of two petal-like lateral plates, each with a vesical denticulated lobe (‘ vdl’ in Figs 30 H, 34 B, C, 38 A, B). This clade is also supported by two homoplasies: males with rows of frontal setae anteriorly divergent, and basiphallus with a dorsal longitudinal keel. Dodge (1965 a) defined the monospecific genus Carinoclypeus by the presence of a ‘ carinate clypeus’. A slightly modified wording for this character state was used by Pape (1996), who diagnosed Carinoclypeus by the presence of a ‘ facial plate with distinct median carina in full length’, which here corresponds to the carina parallel in full length to frontogenal suture. Here, no other character states support this genus, which remains defined only by the autapomorphic presence of a median carina on the facial plate, which supports Dodge’s (1965 a) and Pape’s (1996) diagnoses. Udamopyga (s. s.) is recovered as monophyletic, but it is only supported by one autapomorphy: facial ridge with dense and short setosity on lower 0.50 (Fig. 46 D). Based on the strong branch support of the genus Udamopyga (including Carinoclypeus), and its numerous autapomorphies, we suggest Carinoclypeus as a new junior synonym of Udamopyga. Consequently, we provide a new diagnosis for Udamopyga inclusive of Carinoclypeus, which is maintained as a subgenus, new status.	en	Buenaventura, Eliana, Pape, Thomas (2018): Phylogeny, evolution and male terminalia functionality of Sarcophaginae (Diptera: Sarcophagidae). Zoological Journal of the Linnean Society 183 (4): 808-906, DOI: 10.1093/zoolinnean/zlx070, URL: https://academic.oup.com/zoolinnean/article/183/4/808/4757488
814387FFFFFAFFB104B93F485783FD5E.taxon	description	Within the Peckiamyia clade a sister-group relationship was found between the monophyletic Sinopiella and the remaining genera, arranged in clade 74. The genus Sinopiella is represented in our analysis by its two known species, and it emerges as monophyletic (clade 73 in Fig. 2 B). While all other genera of the Peckiamyia clade have hillae, the lateral styli in the genus Sinopiella are simple and exhibit no modifications. The monophyly of this genus received strong branch support, and its eight autapomorphies are all in the male terminalia: (1) postgonite slightly swollen, (2) postgonite enlarged, (3) pregonite dorso-ventrally flattened and concave, (4) paraphallus humped postero-distally (Fig. 21 D), (5) vesica three-lobed with a proximal section undivided and lobe-shaped (Fig. 21 F), (6) vesical lateral arms elongated with rounded apex (Fig. 21 F), (7) juxta deeply recessed within the phallic tube (Fig. 21 D – F) and (8) juxta squared with ventral margin pointed (Figs 5 G, 21 F). In the description of this genus, Lopes & Tibana (1982) suggested a close relationship with Peckiamyia based on the short phallus, which is supported by our results. In the same publication, these authors also suggested a relationship between Sinopiella and Retrocitomyia due to both genera sharing enlarged pregonites. Although both these genera are closely related as members of the Peckiamyia clade, this sister-group relationship is not recovered in our phylogeny, as the enlarged pregonites were observed only in Retrocitomyia, while Sinopiella has normal-sized pregonites and enlarged postgonites. Kutty et al. (2010) found strong support for a sister-group relationship between Sinopiella rotunda (Lopes & Ferraz, 1991) and Lepidodexia (Notochaeta) sp., but here all species of Lepidodexia form a single clade not closely related to Sinopiella. Finally, the three character states (male mid-femur with ctenidium of rounded spines, wing with third costal sector bare ventrally and three conducting styli) listed by Pape (1996) are not diagnostic for this genus. Clade 74 received weak JK value and is supported by the following autapomorphic character states: (1) hillae directed distally (Figs 16 F, G, 35 C, E), (2) hillae paddle-like (Figs 16 F, 35 C – E), (3) only apex of hillae attached to the inner paraphallic wall (Figs 16 F) and (4) juxta squared with anterior margin even (Figs 16 H, 35 F, 36 E). The presence of proximal expansions of the lateral styli or hillae in clade 74 is homoplasious in our analysis and appears to have evolved in the ancestor of the Tricharaea grade or earlier, becoming reduced in clade 38, and reappearing in clade 74. Generally, the hillae are visible (Fig. 16 F – H) in lateral view in Duckemyia, while in Peckiamyia, Retrocitomyia and Tapacura they remain hidden by the lateral wall of the distiphallus. In some species of the last three genera, the hillae are distally attached to the inner wall of the juxta, leaving two low swellings that are visible in dorsal view (arrows in Fig. 28 A). Clade 74 is also supported by the presence of a three-lobed vesica, whose proximal section is undivided and arch-shaped in Duckemyia (Fig. 16 F), Retrocitomyia (Fig. 10 G) and Tapacura (Fig. 36 B), while in Peckiamyia this section has a shallow proximal division giving two joined lobes (Fig. 35 D, E). Tapacura is reconstructed as a monophyletic taxon with weak JK value but supported by two autapomorphies: (1) vesical lateral arms disc-shaped (Fig. 36 A, B) and (2) juxta squared with anterior margin even and flat (Fig. 36 E). This genus has very small and distinctive male genitalia, which may carry informative characters for supporting its monophyly. Species of Tapacura have lateral plate-like structures completely fused to the paraphallic wall and with a distal cleft (Fig. 36 A – C, E). The homology of these structures is uncertain. These plate-like structures are in a similar position than the paraphallic blinkers. However, they lack the landmark for delimiting these blinkers, which is a desclerotized strip between them and the ventral margin of the paraphallus. Also, the lateral plates of Tapacura are completely sclerotized, while the paraphallic blinkers are semi-sclerotized. A sister-group relationship between Retrocitomyia (excluding Retrocitomyia argentina Lopes, 1988) and (Duckemyia + Peckiamyia) was recovered in our analysis (clade 76 in Fig. 2 B). Clade 76 is supported by two uniquely derived apomorphies: (1) cercal prong S-shaped with uni- or bilobed apex (Fig. 40 H) and (2) postgonite directed perpendicular to body axis. The monophyly of Retrocitomyia (excluding R. argentina) is strongly supported in our analysis by four autapomorphies: (1) cercal prong bilobed with a blunt tip (see arrows in Fig. 47 A), (2) cercal prong without dorso-medial setae, (3) vesical lateral arms paddle-like with a hook-shaped apex and (4) juxta squared with anterior margin even, undulated dorso-ventrally or with a medial folding (Figs 10 G, H, 28 A). The two tips of the bilobed cercal prong might be more developed in some Retrocitomyia species than in others. Lopes (1983) assigned Retrocitomyia, together with Chlorosarcophaga and Dexomyophora (both included in Lepidodexia [s. l.] by Pape [1996]), and Udamopyga, to the subtribe Udamopygina based on various features of the cephaloskeleton of the first-instar larva, a concavity in ST 8 of the female, the presence of ‘ large lateral plates’ on the distiphallus and the absence of a vesica. Our results did not support a relationship between Retrocitomyia, Lepidodexia and Udamopyga, and each of these genera emerged within separate, distantly related clades. Also, neither of the diagnostic character states proposed by Lopes (1983) in the description of Retrocitomyia nor those suggested by Pape (1996) emerged as autapomorphic for this genus. However, when used in combination, those character states will still be useful for diagnosing this genus. The sister-group relationship between Duckemyia and Peckiamyia has moderate branch support and is supported by three autapomorphies: (1) facial ridge with dense setosity on lower 0.85 (Fig. 46 B), (2) cercal prong bilobed with a pointed tip (Fig. 40 H) and (3) juxta squared, with anterior margin even, flat or slightly concave (Figs 16 F, 35 E). Of the two genera, only Peckiamyia is supported by multiple autapomorphies of the male terminalia and other body parts as follows: (1) postgenal setulae much longer than genal setulae (Fig. 47 B), (2) surstylus with a proximal lobe-shaped expansion, (3) surstylus with stubby setae on proximal half, (4) pregonite with strong proximal setae, (5) vesica three-lobed, whose proximal section has a shallow proximal division giving two joined lobes (Fig. 35 D, E) and (6) vesical lateral arms trapezoid (Fig. 35 C, E). Duckemyia shows one autapomorphy: vesical lateral arms ribbon-like (Fig. 16 G). Dodge (1966) identified similarities in external characters between Peckia and Peckiamyia, but he also mentioned Peckiamyia as having ‘ anomalous genitalia’ obscuring its affinities. A close relationship between Peckia and Peckiamyia has not been supported in subsequent studies (Piwczyński et al., 2014; Buenaventura & Pape, 2015), nor in the present study. A comparison of features of Duckemyia latifrons Kano & Lopes, 1969 to those of potentially close generic relatives with proclinate fronto-orbital setae in males was provided by Kano & Lopes (1969), who erected a separate genus for this species. The proclinate fronto-orbital setae in males were here found to be a homoplasious character state. In the same publication, these authors also noted the bifurcated cercal prong (Fig. 40 H) in Duckemyia and Peckiamyia, which is also shared with Retrocitomyia.	en	Buenaventura, Eliana, Pape, Thomas (2018): Phylogeny, evolution and male terminalia functionality of Sarcophaginae (Diptera: Sarcophagidae). Zoological Journal of the Linnean Society 183 (4): 808-906, DOI: 10.1093/zoolinnean/zlx070, URL: https://academic.oup.com/zoolinnean/article/183/4/808/4757488
814387FFFFF9FFBF07133EB151C7FCB8.taxon	description	Our analysis recovered a monophyletic genus Boettcheria as sister to the clade (Microcerella + (Austrophyto + R. argentina )). Based mostly on male terminalia characters, Roback (1954) placed Boettcheria close to Sarcodexiopsis and Tripanurga and included these genera in the subtribe Boettcheriina. Lopes (1983) placed this subtribe within Sarcophagini, but he restricted Boettcheriina to species of Boettcheria, and in the same publication he suggested a possible relationship between Boettcheriina and Microcerellini, this last tribe containing species with ‘ bare or pubescent arista’. In a subsequent publication, Lopes (1989) described Austrophyto as a monospecific genus and placed it into the tribe Microcerellini. Pape (1990) synonymized all the generic names included in the Microcerellini of Lopes (1983) under Microcerella, excluding only Cryptosarcophila Townsend (transferred to Lepidodexia as a subgenus) and Austrophyto. Pape (1994) found Boettcheria as sister to Emdenimyia, a relationship supported by the configuration of postero-ventral setae on the trochanter (see discussion of the Blaesoxipha clade), but he did not include Austrophyto or any species of Microcerella. Based on molecular data, Kutty et al. (2010) found Boettcheria cimbicis (Townsend, 1892) as part of a trichotomy with Engelimyia inops (Walker, 1849) and Tricharaea femoralis (Schiner, 1868); Stamper et al. (2012) recovered a monophyletic Boettcheria as sister to T. importuna, and Piwczyński et al. (2014) recovered a sister-group relationship between a monophyletic Boettcheria and E. inops. Thus, molecular data do not yet converge in their phylogenetic estimations with regard to the position of Boettcheria, while the morphological data of Giroux et al. (2010) coincide with ours in placing Boettcheria and Microcerella as closely related taxa. Lopes (1950) revised the species of Boettcheria and provided a definition for this genus, where he highlighted the characteristic shape of the male ST 5 and the very large vesica. Pape (1989 b) redefined this genus and proposed a diagnosis including four character states. In a subsequent revisionary work of the Nearctic species of Boettcheria, Dahlem & Downes (1996) provided a generic definition based on three character states. Pape (1996) proposed a diagnosis for Boettcheria, in which he included some of his own character states (Pape, 1989 b) and also those of Dahlem & Downes (1996). Here we included all of Pape’s (1996) diagnostic character states, of which two were reinterpreted and combined into one character state. Our analysis resulted in five autapomorphies supporting this genus. Pape’s character state of the modified setae on the male hind trochanter is separated into two character states, with male hind trochanter with a postero-ventral brush-like clump of short, stubby setae distally (position as no. 1 in Fig. 45) coming out as an autapomorphy for Boettcheria. The remaining four autapomorphies are: (1) six or more frontal setae below posterior limit of the lunule, (2) male abdominal T 5 higher than other abdominal tergites, (3) vesica convoluted (Fig. 30 F) and (4) juxta squared with proximal corners slightly elongated (Fig. 30 F). Character state 3 may be seen as a simplified way of describing the most complex structure in the male terminalia of species of Boettcheria. The vesica in this genus has been previously described as ‘ trilobed’ (Dahlem & Downes, 1996) or ‘ with more than three lobes’ (Giroux et al., 2010); however, any subdivision into lobes or a more detailed definition of this structure would require a homology assessment based on a more inclusive sample of species, which is not the scope of our study. Additional characters with potential phylogenetic content are (1) the unusually larger membranous area between the epandrium and the proximal margin of the surstylus and (2) the L-shaped surstylus in most species of this genus. The sister-group relationship between (Austrophyto + R. argentina) and Microcerella received strong JK value and is supported by two autapomorphies: (1) male hind trochanter with a pad of short setae covering almost the entire posterior surface (position as no. 3 in Fig. 45) and (2) phallus with a paler ventral area between basi- and distiphallus (arrow in Figs 38 E, 47 G). Mulieri (2017) revised Austrophyto and provided a definition for this genus, where he highlighted several features, most of them found in many other genera in Sarcophaginae, but also including the (1) postgonite with two long setae, (2) distiphallus with a swollen, desclerotized ventral area proximal to vesica, (3) vesica short and weakly sclerotized, with a microserrated margin and (4) juxta scarcely developed, with apico-lateral membranous lobes and a medial sclerotization (= medial juxtal sclerite) between them. Character state 1 was included here in its original form, while state 2 was included as homologized with paler ventral area between basi- and distiphallus being swollen in Austrophyto and R. argentina, state 3 was included as vesica with a proximal desclerotized, microserrated and bilobed section (‘ vbs’ in Fig. 38 E, G) and state 4 was divided into median juxtal sclerite (‘ mjs’ in Fig. 38 F, G) and juxta as two apico-lateral membranous lobes (‘ jl’ in Fig. 38 F – H). Mulieri (2017) also highlighted the distiphallus with ‘ strongly developed harpes’, which were not included here due to lack of material. However, the harpes in this genus are conspicuous with a shape not observed in other genera of Sarcophaginae. Mulieri (2017) also compared the reduced juxta of Austrophyto with that of Boettcheria; however, we do not find support for the latter genus having a juxta reduced nor morphologically similar to that of Austrophyto. Some additional comments by Mulieri (2017) on the possible phylogenetic relatedness of Austrophyto to Boettcheria and Microcerella were not endorsed by phylogenetically informative evidence and are considered unsupported. Our analyses reconstructed the monophylum of (Austrophyto + R. argentina), which received strong JK value and is supported by five autapomorphies: (1) postgonite with two long setae (Fig. 38 D), (2) paler ventral area between basi- and distiphallus swollen (arrow in Fig. 38 E), (3) vesica with a proximal desclerotized, microserrated and bilobed section (‘ vbs’ in Fig. 38 E, G), (4) median juxtal sclerite (‘ mjs’ in Fig. 38 F, G) and (5) juxta as two apico-lateral membranous lobes (‘ jl’ in Fig. 38 F, G). The affinity of R. argentina was uncertain also for Lopes (1988 b), who assigned it provisionally to Retrocitomyia in spite of the absence of terminalia features typical of that genus. Based on our phylogeny and morphological examinations, we propose to include R. argentina in Austrophyto, with Austrophyto argentina (Lopes, 1988) as a new combination. Previous definitions for Microcerella (Macquart, 1851; Hall, 1937; Lopes, 1983; Pape, 1996) were considered as ‘ skewed’, ‘ based on highly homoplastic characters’, and as considering only ‘ few and unuseful character states’ (Mulieri et al., 2015). However, subsequent definitions for this genus included some character states such as ‘ male without orbital proclinate setae’ (Mulieri et al., 2015), which is not diagnostic for this genus, since it is found in at least 37 genera of Sarcophaginae. Outlining a definition for this and other Sarcophaginae genera compels researchers to use homoplasies, which are abundant in the subfamily, as already reported (Giroux et al., 2010). This overwhelming level of homoplasy could have resulted from multiple specializations giving morphologies that retain few clues to their phylogenetic history. In the description of the genus Microcerella, Macquart (1851) used the bare arista to define this taxon, which was also included in definitions proposed by subsequent authors (Hall, 1937; Lopes, 1983; Pape, 1990, 1996). Pape (1990) defined Microcerella by the following character states: (1) eyes green, (2) syntergosternite 7 + 8 black, (3) hypandrium swollen at level of pregonite, (4) postgena with at least some black setae close to genal suture, and he also pointed to the (5) syntergosternite 7 + 8 dark brown to black / epandrium red. Pape (1996) added two other character states: (6) strong parafacial setae, and (7) arista almost bare. Mulieri et al. (2015) included (8) three strong postsutural dorso-central setae, (9) rigid connection between basi- and distiphallus, fused anteriorly with an incomplete hinge on posterior part, and (10) phallus with a paler anterior (= ventral) area between disti- and basiphallus. From these ten character states, 1, 3 and 5 came out as autapomorphic for this genus in the present study, while character state 2 was also found in Boettcheria and 10 was also found in Austrophyto, character states 7 and 9 were autapomorphic for the entire Microcerella clade and states 4, 5, 6 and 8 were homoplasious. Besides character states 1, 3 and 5, the monophyly of Microcerella was also supported by the paler and flat ventral area between basi- and distiphallus (Fig. 47 G) (swollen in Austrophyto), and the juxta campanulated to oval (Figs 29 F, G, 47 G).	en	Buenaventura, Eliana, Pape, Thomas (2018): Phylogeny, evolution and male terminalia functionality of Sarcophaginae (Diptera: Sarcophagidae). Zoological Journal of the Linnean Society 183 (4): 808-906, DOI: 10.1093/zoolinnean/zlx070, URL: https://academic.oup.com/zoolinnean/article/183/4/808/4757488
814387FFFFF7FFBD07133E07551AFAB1.taxon	description	The species currently assigned to the genus Lepidodexia possess similarities in the phallic morphology, although their diversity in external morphology is remarkable (Lopes, 1951, 1979, 1984, 1985, 1991, 1992). Some of these similarities were noticed by Roback (1954), who considered Camptops Aldrich, Chloronesia Townsend, Harpagopyga Aldrich, Johnsonia and Notochaeta to be phylogenetically close and placed them in the Johnsonia group. Roback (1954) also included Argoravinia, Emblemasoma and Helicobia in this group. Similarly, Lopes (1979, 1984) proposed the tribe Johnsoniini, where he included all the subgenera currently assigned to Lepidodexia plus some species currently in the genera Archimimus and Emdenimyia. The Johnsoniini of Lopes share character states of the head chaetotaxy, female terminalia, labrum of the first-instar larva and male terminalia structures such as the ‘ spinous lobe of the vesica’ (Lopes, 1979, 1984). Lopes (1983) also included Malacophagula and Rafaelia in the tribe Johnsoniini. Almost all the genera belonging to the Johnsoniini of Lopes were synonymized under Lepidodexia by Pape (1995, 1996), being characterized by the vesica bearing a proximal spinous lobe, and only excluding species of Archimimus and Emdenimyia, to produce a Lepidodexia (sensu lato) containing 29 subgenera. Many of these are monospecific: Chloronesia, Cryptosarcophila, Halliosca, Neophytodes Townsend, Orodexia Townsend, Paramintho Brauer & Bergenstamm, Petriana Lopes and Stenopygopsis Townsend; others include only a few species, for example Abacantha Hall, Dexomyophora, Eufletcherimyia Townsend, Geijskesia Lopes, Hallina and Travassosisca Lopes, while only six subgenera have numerous species, i. e. Chlorosarcophaga, Harpagopyga, Johnsonia, Lepidodexia, Neophyto and Notochaeta. Neither the genus Lepidodexia nor any of its subgenera have been recently revised. Only three phylogenetic studies have included species of this genus (Lopes, 1984; Giroux et al., 2010; Kutty et al., 2010), and only one of these (Giroux et al., 2010) found Lepidodexia as monophyletic, although this clade was supported only by homoplasies. Thus, the monophyly of Lepidodexia and its subgenera had not been consistently tested, and there is no phylogenetic hypothesis for relationships within this genus. In the present study, we included representative species of only six subgenera, i. e. Lepidodexia (Chlorosarcophaga), L. (Dexomyophora), L. (Hallina), L. (Halliosca), L. (Neophyto) and L. (Notochaeta), of which all represented by more than one species emerged as monophyletic within a paraphyletic genus Lepidodexia (Fig. 2 B). The only species of Halliosca emerges near the base of the Lepidodexia clade as it lacks the two autapomorphies that support this clade: (1) the presence of a hinge between the proximal and distal parts of the harpes [fused in Halliosca (Fig. 31 E, F)], and (2) juxta angled [arching in Halliosca (Fig. 31 E)]. Halliosca shows several character states shared with the genus Lipoptilocnema, including male abdominal ST 5 with two pointed black cuticular processes on the angle of the V-shaped cleft, margin of surstylus overlapping the hinge between epandrium and surstylus, and cercal prong bent at mid-length, and with a proximal tuft of long black setae (identical to those of Lipoptilocnema). As outlined above, Pape (1996) proposed a broadened concept of Lepidodexia (sensu lato) containing 29 subgenera, one of these being Halliosca. The strong support found for a sister-group relationship between Emblemasoma and Lepidodexia (exclusive of Halliosca) leaves two options: to exclude Halliosca as subgenus from the genus Lepidodexia (Pape, 1996), or to broaden the definition of the latter to include also Archimimus and Emblemasoma. We are here resurrecting Halliosca as a valid genus, new status. Pape (1996) diagnosed Lepidodexia with three character states: (1) postalar wall bare, (2) distiphallus with juxta angled relative to the phallic tube (Fig. 32 A) and (3) distiphallus with a spinous lobe proximal to the vesica (no. 1 in Figs 31 D, 32 C, F). Character state 1 is not particularly diagnostic for Lepidodexia, since it is shared only by the subgenera L. (Neophyto) and L. (Notochaeta). As mentioned above, character state 2 emerged as autapomorphic for clade 90, as it is shared by all members of the Lepidodexia clade except Halliosca (Fig. 31 E – G). Character state 3, originally described by Lopes (1979, 1984), is autapomorphic for clade 92, which consists of all subgenera of Lepidodexia (including Archimimus), together with three uniquely derived synapomorphies: (1) arista almost twice as long as first flagellomere (Fig. 44 F), (2) male abdominal ST 5 with a rounded expansion taking up the entire posterior half (Fig. 44 E), (3) vesica bipartite with a C-shaped medial section (no. 3 in Fig. 32 C) and a convex sclerotized distal section (no. 2 in Fig. 32 C). A comparison of the proximal spinous lobe of the vesica in various subgenera of Lepidodexia shows that this feature can be homologized across the genus (red structure in Fig. 33). A monophyletic Lepidodexia can be attained by either raising all subgenera to valid genera, lumping all species into a Lepidodexia (sensu lato), or a combination of the two. Following the last option, and in order to attain a monophyletic Lepidodexia, we choose to include Archimimus in this genus, as a subgenus, new status, and exclude Halliosca and give it the new status as a valid genus. This newly circumscribed Lepidodexia (including Archimimus) received strong branch support and is supported by the conspicuous proximal spinous lobe of the vesica plus the above-mentioned autapomorphies. The monophyly of and relationships between the subgenera of Lepidodexia are partially supported. Thus, L. (Dexomyophora) is supported by a facial ridge with dense setosity on lower 0.70 (Fig. 46 C), while L. (Hallina), L. (Neophyto) and L. (Notochaeta) are only supported by homoplasies. Three out of five of the currently recognized species of Archimimus (sensu Pape, 1996) are included in the present study, and they formed a strongly supported monophyletic group that emerged as sister to L. (Neophyto). The monophyly of L. (Archimimus) is supported by three autapomorphies: (1) pregonite distally spatulated, (2) median stylus truncated (Fig. 9 G) and (3) median stylus with no opening (Fig. 9 G). Only five genera and one subgenus of Sarcophaginae have the median stylus strongly modified into an apparently non-conducting stylus or entirely reduced. These are L. (Archimimus), Chrysagria, Helicobia, Lipoptilocnema, Peckia and Sarcophaga, and all are characterized by different acrophallic configurations. Lepidodexia (Archimimus) and Lipoptilocnema have both a median stylus and a capitis, but the median stylus is not tubular (Figs 9 G, 24 B – I); Chrysagria has a short capitis and an entirely reduced median stylus (Fig. 11 C); Helicobia and Sarcophaga have an elongated capitis and an entirely reduced median stylus (Fig. 37 C, G); and Peckia has no trace of either a median stylus or a capitis (Figs 12 H, 13 H). The sister-group relationship between L. (Archimimus) and L. (Neophyto) is supported by one autapomorphy: distance between occiput and antennal base longer than distance between occiput and vibrissal angle. Roback (1954) included Emblemasoma in the Johnsonia group, and considered it to be closely related to Helicobia and Johnsonia (= Lepidodexia, in part) due to structural similarities in the male terminalia. Emblemasoma was considered as part of the tribe Sarcophagini by Lopes (1969 a), but Lopes (1983) later erected the tribe Emblemasomatini for Emblemasoma and Pessoamyia Lopes. Our results support these assumptions, since Emblemasoma is closely related to Lepidodexia, as suggested by Roback (1954), and species originally in Emblemasoma and Pessoamyia constitute a monophylum, as indicated by Lopes (1969 a). Lopes (1971) defined Emblemasoma and Pessoamyia by the presence of an inflated prosternum. Pape (1996) synonymized these two genera and expanded the definition of Emblemasoma, which he diagnosed as follows: (1) prosternum enlarged, (2) male mid-femur with a ctenidium of rounded spines (circular cross section) and (3) male cercus distally swollen and with a blunt tip (Fig. 44 G). Here, the monophyly of Emblemasoma was tested for the first time, and it is supported by seven autapomorphies, mostly from non-terminalia characters. These include character states 1 and 3 of Pape (1996), plus facial plate almost equibroad along its entire length (Fig. 44 A), parafacial plate widest at level of lunule (Fig. 44 B), palpus with long setae (Fig. 44 B), male mid-femur with 1 – 4 antero-dorsal setae at mid-length and vesica composed of two leaf-shaped lobes (Fig. 23 A – E).	en	Buenaventura, Eliana, Pape, Thomas (2018): Phylogeny, evolution and male terminalia functionality of Sarcophaginae (Diptera: Sarcophagidae). Zoological Journal of the Linnean Society 183 (4): 808-906, DOI: 10.1093/zoolinnean/zlx070, URL: https://academic.oup.com/zoolinnean/article/183/4/808/4757488
814387FFFFF5FFBB048B380C5782FEEF.taxon	description	The monophyly of Helicobia has been supported by morphological (Giroux et al., 2010; Buenaventura & Pape, 2015) and molecular studies (Kutty et al., 2010; Stamper et al., 2012; Piwczyński et al., 2014; Buenaventura & Pape, 2017) and it is also strongly supported by our results. Giroux et al. (2010) reduced Helicobia to a subgenus of Sarcophaga, but this was rejected by subsequent studies (Kutty et al., 2010; Stamper et al., 2012; Piwczyński et al., 2014; Buenaventura & Pape, 2015, 2017), as well as by our results. A single autapomorphy supported this genus: the male hind trochanter with a pad of short setae medially and with a strong seta near its posterior margin (position as no. 7 in Fig. 45). Of the seven apomorphies that supported this taxon in Giroux et al. ’ s (2010) phylogeny, two – posterior and postero-ventral setae in the male hind tibia unmodified and dorsal proximal part of wing vein R 1 setulose – were included here, and found not to be uniquely derived in this genus but shared with at least 15 other genera. Another homoplasious character state supporting Helicobia is a parafacial plate with strong setae. Similarly, of the six character states defining Helicobia in Buenaventura & Pape’s (2015) study, five are included here but are not recovered as autapomorphic for this genus. Two of them (ocellar setae strong, vertical setae strong) do not define Helicobia in our study, while the three remaining ones correspond to configurations of the vesica that are here reinterpreted. Female T 6 with a mid-dorsal desclerotized, fine strip or narrow membranous longitudinal cleft was not included in the present study, due to scarce female data for other Sarcophaginae genera. Despite the homoplastic condition of character states in the present study as well as those of Pape (1996), Giroux et al. (2010) and Buenaventura & Pape (2015), we use a combination of these to define Helicobia. The clade (Peckia + (Lipoptilocnema + Sarcophaga )) is supported by one autapomorphy: cercal prong with a subapical saddle-shaped concavity followed by a hump. This clade was also supported by two homoplasies: (1) postgenal setulae white or yellow, and (2) one presutural dorso-central seta. Peckia and Sarcophaga also share an inner margin of male abdominal ST 5 cleft with a large medial pad of long hair-like setulae, or strong and short setae. This setosity pattern is absent in Lipoptilocnema, which instead has two pointed black cuticular processes on the angle of the V-shaped cleft of the male abdominal ST 5. Buenaventura & Pape (2015) included all currently recognized species of Peckia (sensu Buenaventura & Pape, 2013) and provided an extensive discussion on the historical definitions and concepts of this genus by especially Robineau-Desvoidy (Robineau-Desvoidy, 1830), Lopes (1941 a, 1943, 1958, 1969 a, 1983), Roback (1954) and Pape (1996). Two synapomorphies supported the monophyly of Peckia in Buenaventura & Pape (2015): (1) presence of a fringe of long, hair-like setulae along outer margin, extending to – or almost to – the posterior corner of the lower calypter, and (2) reduction of the capitis. These character states, plus paraphallus wider than long (Fig. 13 E, F) also support Peckia in our analysis. The paraphallic tube in Peckia is mostly reduced (except in the subgenus Pattonella Enderlein, Fig. 12 F), consisting almost only of a sclerotized strip in the proximal part of the distiphallus, whereas the juxta is generally large and complex, particularly in the subgenera Pattonella (Fig. 12 F), Peckia (Fig. 13 F – H) and Sarcodexia (Fig. 35 B). For example, the juxta in the subgenus Sarcodexia has one basal and two distal juxtal horns (‘ bjh’ and ‘ djh’ in Figs 13 I, J, 35 A, B). The genus Peckia is also supported by three homoplasies, including the loss of harpes. All groups in basal positions with regard to clade 80 have a distiphallus with no harpes. According to the optimization of this character in our phylogeny, the harpes are considered as primarily absent in the Tricharaea and Dexosarcophaga grades, and the clades Oxysarcodexia, Argoravinia, Blaesoxipha, Engelimyia, Udamopyga and Peckiamyia, but present in clades Microcerella, Lepidodexia and Sarcophaga, while in the genus Peckia they are secondarily lost, which may constitute a reduction uniquely derived in this genus. The clade (Lipoptilocnema + Sarcophaga) received high JK value and is supported by three uniquely derived character states: margins of surstylus slightly folded or protruding outwards (‘ sr’ in Fig. 44 C, D), paraphallic dorsal wall with a longitudinal desclerotized strip with a shallow or deep depression (Figs 24 J, 37 H) and presence of paraphallic proximal expansions (‘ ppe’ in Figs 24 C, 37 E, I). This clade was also supported by three homoplasies: male with rows of frontal setae divergent anteriorly, cercus with proximal tuft of long, black, hair-like setulae and harpes protruding dorso-medially over the base of the lateral styli (Fig. 37 E, I). Buenaventura & Pape (2015) interpreted the acrophallic structures of the genera of the Sarcophaga clade, such as the reduced median stylus and the elongated capitis, in the same way as here, but some additional character states included in the present analysis resulted in a change in relationships among these genera. Thus, some character states such as the subapical saddle-shaped concavity of the cercal prong followed by a subapical hump support the clade (Peckia + (Lipoptilocnema + Sarcophaga )). Also, the slightly folded or outwards protruding margins of surstylus, the presence of a paraphallic desclerotized strip and the presence of paraphallic proximal expansions support the clade (Lipoptilocnema + Sarcophaga).	en	Buenaventura, Eliana, Pape, Thomas (2018): Phylogeny, evolution and male terminalia functionality of Sarcophaginae (Diptera: Sarcophagidae). Zoological Journal of the Linnean Society 183 (4): 808-906, DOI: 10.1093/zoolinnean/zlx070, URL: https://academic.oup.com/zoolinnean/article/183/4/808/4757488
