Argoravinia, Townsend, 1917

Argoravinia View in CoL clade

This clade is composed of four genera arranged in the topology ( Malacophagula + Rafaelia ) + ( Argoravinia + Malacophagomyia [including Dodgeisca ]). Most of these nodes received aBS and moderate to strong JK supports in our analysis ( Fig. 2A View Figure 2 ). The Argoravinia clade is supported by five autapomorphies: (1) posterior margin of ST5 very widely V-shaped with an obtuse inner angle ( Fig. 42G View Figure 42 ), (2) paraphallus dorso-distally rounded ( Figs 14C View Figure 14 , 15D View Figure 15 , 17A, F View Figure 17 ), (3) vesica broad and flat ( Figs 15F View Figure 15 , 17A, B, D View Figure 17 ), (4) hillae directed latero-ventrally ( Figs 15A View Figure 15 , 16C View Figure 16 , 17A View Figure 17 ) and (5) hillae not touching the inner paraphallic wall ( Fig. 15E View Figure 15 ).

The clade ( Malacophagula + Rafaelia ) is supported by four homoplasies: parafacial plate with strong setae ( Fig. 42F View Figure 42 ), 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 14B View Figure 14 , 17G View Figure 17 ). 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. 42F View Figure 42 ), first flagellomere shortened ( Fig. 42F View Figure 42 ), lunule widened, postgena swollen ( Fig. 42F View Figure 42 ) and lower calypter rounded ( Fig. 43A View Figure 43 ).

One autapomorphy and three homoplasies supported the monophyly of Rafaelia , which received moderate branch support ( Fig. 2B View Figure 2 ). 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 20F View Figure 20 , 40C, D View Figure 40 ), 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 15A–C View Figure 15 , 16C View Figure 16 , 17A View Figure 17 ) and (3) median stylus greatly elongated ( Figs 15B, E View Figure 15 , 16D View Figure 16 , 17A, B, D View Figure 17 ).

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. 15B–F View Figure 15 ), (4) capitis as a smooth, rounded lobe, proximally swollen, (5) median stylus S-shaped ( Fig. 15B, E View Figure 15 ) and (6) juxta very small to vestigial ( Fig. 15E View Figure 15 ). Some of these character states were previously included in the generic diagnoses for Argoravinia ( Lopes, 1976a; Pape, 1990, 1996; Carvalho-Filho & Esposito, 2012). For example, Lopes (1976a) 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. 2A View Figure 2 ). 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 T6, 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 (1969a, 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. 17A, B, D View Figure 17 ) and (7) juxta arching over the lateral styli ( Fig. 17A, B, D View Figure 17 ). Interestingly, the most remarkable character states (6 and 7) are shared with the species Dodgeisca paramerata Rohdendorf, 1971 ( Fig. 16C, D View Figure 16 ), 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 ST4 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 16C, D View Figure 16 , 17A, B, D View Figure 17 ), 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 16C–E View Figure 16 , 17A, B View Figure 17 ), and their ‘harpes’ with the paraphallic lateral expansions ( Figs 16C View Figure 16 , 17A View Figure 17 ). 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 16D View Figure 16 , 17B View Figure 17 ), 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.