Sarcophaga

Sarcophaga View in CoL clade

The Sarcophaga View in CoL clade (clade 101 in Fig. 2B View Figure 2 ) is formed by Chrysagria View in CoL as sister to ( Helicobia View in CoL + ( Peckia View in CoL + ( Lipoptilocnema View in CoL + Sarcophaga View in CoL ))) and all genera were reconstructed as monophyletic. The entire clade has weak support and half of its internal branches have high branch supports. Apart from Chrysagria View in CoL and Lipoptilocnema View in CoL , all genera of the Sarcophaga View in CoL clade had been included in previous phylogenetic analyses. Giroux et al. (2010) found Sarcophaga View in CoL as paraphyletic with regard to Helicobia View in CoL , and Peckia View in CoL as the sister group of ( Sarcodexia View in CoL + Titanogrypa View in CoL ). Few species of Helicobia View in CoL and Peckia View in CoL , and representatives of 31 subgenera of Sarcophaga View in CoL were included. Kutty et al. (2010) found a monophyletic Helicobia View in CoL only distantly related to Sarcophaga View in CoL and to a paraphyletic Peckia View in CoL . Stamper et al. (2012) found Helicobia View in CoL as sister to ( Peckia View in CoL + Sarcophaga View in CoL ), whereas Piwczyński et al. (2014) found ( Peckia View in CoL [including Villegasia View in CoL ] + Sarcophaga View in CoL ) as the sister clade of ( Helicobia View in CoL + (( Boettcheria View in CoL + Engelimyia View in CoL ) + ( Duckemyia View in CoL + Peckiamyia View in CoL ))). Buenaventura & Pape (2015) discussed the monophyly and phylogenetic relationships of four of the five genera included in the present Sarcophaga View in CoL clade. These authors included all currently recognized species of Peckia View in CoL , and the resulting topology, with Peckia View in CoL as sister to ( Lipoptilocnema View in CoL + ( Helicobia View in CoL + Sarcophaga View in CoL )), was generally strongly supported. Buenaventura & Pape (2017) found Helicobia View in CoL as sister to (( Lipoptilocnema View in CoL + Peckia View in CoL ) + Sarcophaga View in CoL ) based on a data set of four molecular markers and species of all biogeographic regions. Differences to the present study are due to the additional male terminalia character states as discussed below.

The Sarcophaga View in CoL clade is well supported. Two autapomorphies define this clade: (1) acrophallus with two styli, and (2) median stylus strongly modified into an apparently non-conducting stylus or entirely reduced (capitis present or not). The first split of the Sarcophaga View in CoL clade shows the genus Chrysagria View in CoL as sister to clade 103, which contains the remaining genera. The genus Chrysagria View in CoL was defined by Lopes & Achoy (1986) by the small ‘apical plate’ (= juxta) and the styli becoming free, among other male and female character states. Pape (1996) also noticed the particular development of the lateral styli in this genus, as one of the diagnostic character states he proposed for this genus was lateral styli long and curved, reaching beyond the apex of the distiphallus. However, this feature is not exclusively found in Chrysagria View in CoL , but is present also in Helicobia View in CoL , Peckia View in CoL and Sarcophaga View in CoL . Our results are not consistent with those of Lopes (1969a, 1983), who included Chrysagria View in CoL and genera like Microcerella View in CoL in the tribe Microcerellini . The three known species of Chrysagria ( Pape, 1996) View in CoL , two of which were included in the present study, form a monophylum receiving strong JK value and supported by two autapomorphies: (1) cercal prong with a median tuft of brown and yellow, medially directed setae, and (2) juxta composed of two elongated and smooth segments ( Fig. 11C View Figure 11 ).

The clade ( Helicobia View in CoL + ( Peckia View in CoL + ( Lipoptilocnema View in CoL + Sarcophaga View in CoL ))) is supported by three autapomorphies: (1) capitis recurved ( Figs 24G View Figure 24 , 37C View Figure 37 ), (2) juxta dome-shaped ( Figs 22I View Figure 22 , 32F View Figure 32 , 37A, D View Figure 37 ) and (3) juxta with juxtal lateral plates (‘jlp’ in Figs 13F View Figure 13 , 22G, I View Figure 22 , 35B View Figure 35 , 37E, F View Figure 37 ). Although the capitis is noticeably developed in Lipoptilocnema View in CoL , Helicobia View in CoL and Sarcophaga View in CoL , it is reduced in Peckia View in CoL . The juxta is generally dome-shaped in this clade; however, in Lipoptilocnema View in CoL it is a membranous expansion covered with sclerotized apical spines ( Mulieri et al., 2016), having a recurved shape, and lacking the juxtal lateral plates.

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 View Figure 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 T6 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 ST5 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 ST5.

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 (1941a, 1943, 1958, 1969a, 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. 13E, F View Figure 13 ) also support Peckia in our analysis. The paraphallic tube in Peckia is mostly reduced (except in the subgenus Pattonella Enderlein , Fig. 12F View Figure 12 ), 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. 12F View Figure 12 ), Peckia ( Fig. 13F–H View Figure 13 ) and Sarcodexia ( Fig. 35B View Figure 35 ). For example, the juxta in the subgenus Sarcodexia has one basal and two distal juxtal horns (‘bjh’ and ‘djh’ in Figs 13I, J View Figure 13 , 35A, B View Figure 35 ). 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. 44C, D View Figure 44 ), paraphallic dorsal wall with a longitudinal desclerotized strip with a shallow or deep depression ( Figs 24J View Figure 24 , 37H View Figure 37 ) and presence of paraphallic proximal expansions (‘ppe’ in Figs 24C View Figure 24 , 37E, I View Figure 37 ). 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. 37E, I View Figure 37 ). 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 ).

Lipoptilocnema View in CoL , represented in this analysis by two species, is defined by four autapomorphies in the male terminalia: (1) proximal margin of surstylus overlapping the hinge between epandrium and surstylus (arrow in Fig. 44C View Figure 44 ), (2) distal part of harpes membranous ( Fig. 24B–D, G View Figure 24 ), (3) juxta recurved ( Fig. 24B, G View Figure 24 ) and (4) juxta triangular with longitudinal keel, laterally membranous, and apically bifid and spinose ( Figs 24J View Figure 24 , 44D View Figure 44 ). The position of this genus within the Sarcophaginae View in CoL was recently analysed by Buenaventura & Pape (2015, 2017), who recovered a monophyletic Lipoptilocnema View in CoL not nested inside any other genus and thereby refuted the proposal of Pape (1996) to include it as a subgenus of Sarcophaga View in CoL . The present phylogeny finds Lipoptilocnema View in CoL as the sister group of Sarcophaga View in CoL as opposed to ( Helicobia View in CoL + Sarcophaga View in CoL ) of Buenaventura & Pape (2015) and ( Lipoptilocnema View in CoL + Peckia View in CoL ) of Buenaventura & Pape (2017). Buenaventura & Pape (2015) found Lipoptilocnema View in CoL as supported by four apomorphies: (1) cercal prong with dorsal surface S-shaped, (2) surstylus with anterior and posterior margin slightly folded, (3) paraphallic apical elongated expansion with apical spines and (4) juxta tongue-shaped, broad proximally and gradually getting narrow to the entire apex. Character state 1 was reinterpreted here and found to be also present in Peckia View in CoL and Sarcophaga View in CoL , while character state 2 was included in its original form and found to be also present in Sarcophaga View in CoL . Character states 3 and 4 were also reinterpreted and homologized to the juxta and median stylus, respectively, in agreement with Mulieri et al. (2016).

Sarcophaga View in CoL is recovered as monophyletic, supported by eight autapomorphies including a medioproximal pad of short setae on the posterior surface of the hind trochanter (position as no. 4 in Fig. 45 View Figure 45 ), a strong seta on postgonite situated distal to middle, paraphallus with a window (‘pw’ in Fig. 37A View Figure 37 ), harpes elbowed in proximal part ( Fig. 37A, E View Figure 37 ) and harpes with an apical process (‘ah’ in Fig. 37F View Figure 37 ). The characteristic paraphallic window of Sarcophaga View in CoL was first described by Whitmore et al. (2013) in a phylogeny of the subgenus Heteronychia . Whitmore et al. (2013) also described the cercal prong of the subgenus Heteronychia with ‘a median, saddle-shaped concavity, or a deep hollowing of the dorsal surface, called a dorsal excavation’, which is also shared by some species of Sarcophaga View in CoL included here. Its autapomorphic condition is only contradicted by its presence in a few species of the subgenus Peckia (Peckia) View in CoL . Sarcophaga View in CoL exclusive of Helicobia View in CoL and Lipoptilocnema View in CoL is a monophyletic taxon, as demonstrated in previous molecular ( Kutty et al., 2010; Stamper et al., 2012; Piwczyński et al., 2014; Buenaventura et al., 2016; Buenaventura & Pape, 2017) and morphological ( Buenaventura & Pape, 2015) studies.