Growing the Rhinophorid Tree: Three New Species and Their Phylogenetic Implications (Diptera: Rhinophoridae)

Simple Summary Rhinophoridae is a family of oestroid flies which was once considered of strictly Palaearctic distribution. This presupposition has been increasingly contradicted in recent years with the discovery and description of many non-Palaearctic rhinophorid species. In this context, the present research article aims to describe three additional species, namely Aporeomyia elaphocera sp. nov., Baniassa pennata sp. nov. and Phyto mambilla sp. nov., from the Oriental and Afrotropical regions. Their genus-level affiliation is supported by a morphology-based phylogeny and these species are described and illustrated, and a comparison with congeners as well as two identification keys are provided. The phylogeny resulting from the addition of these three new species fits well with previous results but retrieves a paraphyletic Phyto Robineau-Desvoidy. Notwithstanding this, the authors were taxonomically conservative, avoiding the erection of a new genus for a single species as well as the lumping of three different genera into a single one: the evidence for doing either of such actions is considered premature and more data are needed. Abstract Three new species of Rhinophoridae (Aporeomyia elaphocera sp. nov., Baniassa pennata sp. nov. from the Oriental Region, and Phyto mambilla sp. nov. from the Afrotropical Region) are described, illustrated and compared with congeners. Genus-level affiliation of the new species is based on a morphology-based phylogeny, preliminarily accepting a paraphyletic Phyto Robineau-Desvoidy awaiting incorporation of molecular data. Keys to the species of the genus Aporeomyia Pape & Shima as well as to the Afrotropical species of the genus Phyto Robineau-Desvoidy are given.

Phyto mambilla sp. nov. is recovered within a clade consisting of other species of Phyto plus Parazamimus and Baniassa, and it emerges as sister to the remaining part of this clade-i.e., ((Phyto adolescens + P. angustifrons) + (Parazamimus Verbeke [19] + Baniassa Kugler)), based on the lack of setosity in the lower half of the parafacial (Matrix S1: 18:0), thus resulting in a paraphyletic Phyto. References: [11,20]. Diagnosis. Male: Medium-sized, slender flies, with relatively large wing. Head profile narrowed ventrally. Prementum and palpus short. Antenna very large and with postpedicel tripartite from base. Arista inserted on anterior lobe. Prosternum bare. Scutellum slightly swollen. Vein M almost straight and without a bend. Veins M 4 and CuA + CuP extended to wing margin. Lower calypter narrow and with median margin diverging from scutellum. Hind tibia with posteroventral apical seta.

Taxonomy
Apomorphies. Male: postpedicel very large and tripartite nearly from base; proclinate orbital setae absent; facial plate deeply sunken; mouthparts strongly reduced; transversal section of sternite 5 almost flat; median extension of surstylus with setae; median process of ventral sclerotization of distiphallus absent. Etymology. The specific epithet, which should be treated as a noun in apposition, is derived from the specific epithet for the red deer, Cervus elaphus Linnaeus, and it refers to the morphology of the postpedicel, tripartite and recalling deer antlers.
Diagnosis. Male postpedicel tripartite from base with the three tips running almost straight (not curved as in A. antennalis Pape & Shima). Base of vein R 4+5 bare dorsally. General ground color dark brown (pale brownish in A. antennalis).   Description. Male (Figures 2 and 3). Length: 4.5-4.7 mm. Color ( Figure 2). Head mostly brown shading into yellowish towards occipital area; frontal vitta black; area between gena and parafacial blackish; scape and pedicel yellowish to light brown; postpedicel and arista blackish-brown; palpus brown. Scutum and scutellum blackish-brown in ground color; scutum covered with barely visible microtomentum except along three thin pre-and postsutural dark vittae (visible in posterodorsal view); postpronotum yellow; thorax in lateral view with two yellow stripes (anterior one descending from postpronotum to fore coxa, across anterior spiracle, proepisternum and proepimeron; posterior one descending from wing base across anterior Description. Male (Figures 2 and 3). Length: 4.5-4.7 mm. Color ( Figure 2). Head mostly brown shading into yellowish towards occipital area; frontal vitta black; area between gena and parafacial blackish; scape and pedicel yellowish to light brown; postpedicel and arista blackish-brown; palpus brown. Scutum and scutellum blackish-brown in ground color; scutum covered with barely visible microtomentum except along three thin pre-and postsutural dark vittae (visible in posterodorsal view); postpronotum yellow; thorax in lateral view with two yellow stripes (anterior one descending from postpronotum to fore coxa, across anterior spiracle, proepisternum and proepimeron; posterior one descending from wing base across anterior half of anepimeron and posterior two-thirds of katepisternum), remaining of pleura brown. Legs dark brown. Upper and lower calypters and wing membrane slightly smoky; tegula brownish and basicosta yellow; wing veins yellow; halter brownish-yellow. Abdomen mostly dark brown in ground color, with anterior portion of tergites 3 and 4 yellowish; tergites 3 and 4 with band of weak, whitish microtomentum on anterior 1/3 (dorsally); terminalia brown.
Head (Figure 2A-D). Head shape receding, roughly triangular in lateral view (i.e., narrowing toward mouthparts). Frons 2.0× as wide as compound eye in dorsal view. Inner vertical seta well developed, 0.8× as long as compound eye height. Outer vertical seta not differentiated. Ocellar triangle with three pairs of short hair-like proclinate setulae (i.e., ocellar seta not differentiated). Frons with 1-4 frontal setae from anterior margin of ocellar triangle and descending at most until middle of pedicel. Fronto-orbital plate with 1 lateroclinate and 3-4 proclinate orbital setae. Upper reclinate orbital setae absent. Parafacial very narrow, as wide as or narrower than width of arista at mid-point. Thorax. Postpronotum with two setae (Figure 2A). Acrostichal setae absent. One presutural and two postsutural dorsocentral setae. One presutural and one postsutural intra-alar setae. First postsutural supra-alar seta absent. Two + one katepisternal setae. Katepimeron and anepimeron bare. One pair of apical scutellar setae, crossed; one pair of subapical scutellar setae. Anatergite with tuft of short setulae below scutellum. Anterior and posterior fringes of posterior spiracle equal in size and emerging from spiracular rim. Legs. Preapical anterodorsal seta of fore tibia about as long as preapical dorsal seta. Mid tibia with one anterodorsal seta. Hind tibia with three preapical dorsal setae. Preapical posteroventral seta of hind tibia about as long as preapical anteroventral seta. Hind tibia with one well-developed anterodorsal seta. Posterodorsal margin of hind coxa bare. Wing (Figure 2A,E). Lower calypter tongue-shaped. Second costal section (CS 2 ) setulose ventrally. Costal spine not differentiated from other costal setulae. Vein R 1 bare. Base of vein R 4+5 bare. Vein M reaching wing margin without bend.
Apomorphies. Posterior lappet of metathoracic spiracle distinctly larger than anterior lappet. Male: Median process of ventral sclerotization of distiphallus interrupted proximally and not connected to ventral plate; surstylus with setae on medial plate. Female: fore tarsus laterally compressed. Etymology. The specific epithet, which should be treated as a Latin adjective, is derived from the Latin "pennatus" meaning "feathered", and it refers to the long, feather-like sclerotizations characterizing the lateral lobes of the distiphallus.
Diagnosis. Parafacial bare in ventral half. Postpronotum with three setae. First postsutural supra-alar seta strong, distinctly longer and thicker than notopleural seta. Bend of vein M not reaching vein R 4+5 so that cell r 4+5 is open at wing margin. Bacilliform sclerite firmly fused to surstylus. Female thorax ground color black.    4× as long as pedicel. Arista with minute trichia shorter than its greatest diameter, and thickened on proximal 1/4. First aristomere very short, not longer than wide. Second aristomere slightly larger than first, not longer than wide. Prementum about twice as long as width at mid length. Palpus apically enlarged.
Diagnosis. Afrotropical species of Phyto likely represent a monophyletic group characterized by a distinctive color pattern with contrasting black and silvery grey microtomentose bands (reminiscent of that of the tachinid genus Trigonospila Pokorny [32]). Lunule with setae. Parafacial bare. First postsutural supra-alar seta present and well developed, longer than notopleural setae. Male: frons very narrow (narrower than width of postpedicel); fronto-orbital plate without proclinate orbital setae.
Apomorphies Etymology. The specific epithet, which should be treated as a noun in apposition, refers to the type locality.
Diagnosis. Male arista with long microtrichia (at most twice the maximum diameter of arista). Parafacial bare in ventral half. Postpronotum with two setae. Bend of vein M not reaching vein R4+5 so that cell r4+5 is open at wing margin. Katepimeron with few hair-like setulae.  Townsend (1916:8).
Diagnosis. Afrotropical species of Phyto likely represent a monophyletic group characterized by a distinctive color pattern with contrasting black and silvery grey microtomentose bands (reminiscent of that of the tachinid genus Trigonospila Pokorny [32]). Lunule with setae. Parafacial bare. First postsutural supra-alar seta present and well developed, longer than notopleural setae. Male: frons very narrow (narrower than width of postpedicel); fronto-orbital plate without proclinate orbital setae.
Apomorphies Etymology. The specific epithet, which should be treated as a noun in apposition, refers to the type locality.
Diagnosis. Male arista with long microtrichia (at most twice the maximum diameter of arista). Parafacial bare in ventral half. Postpronotum with two setae. Bend of vein M not reaching vein R 4+5 so that cell r 4+5 is open at wing margin. Katepimeron with few hair-like setulae.   Description. Male (Figures 8 and 9). Length: 4.3 mm. Color ( Figure 8A-F). Head black in ground color and covered with silvery grey microtomentum; frontal vitta brownish; area between gena and parafacial brownish to orange; scape, pedicel and postpedicel dark brown; palpus orangish. Thorax black in ground color; postpronotum silvery grey microtomentose; prescutum with two broad silvery grey microtomentose vittae; anterior 3/5 of scutum black and posterior 2/5 silvery grey microtomentose; scutellum black; anepisternum and katepisternum silvery grey microtomentose, remaining pleura black. Legs light brown shading to dark brown tarsi. Upper and lower calypters whitish. Wing hyaline; tegula and basicosta brown; veins yellow. Halter whitish. Abdomen brown in ground color with two broad silver bands; syntergite 1 + 2 with brown mid-dorsal depression and silver lateral and ventral parts; tergites 3-4 each with silver band on anterior half, almost divided by brown triangle in the middle; terminalia brown.
Head ( Figure 8A,B,D-F). Frons at narrowest point much narrower than postpedicel, not more than twice as wide as anterior ocellus, 0.072 of head width and 0.15× as wide as compound eye in dorsal view. Outer vertical seta not differentiated. Ocellar setae well developed, proclinate.  Description. Male (Figures 8 and 9). Length: 4.3 mm. Color ( Figure 8A-F). Head black in ground color and covered with silvery grey microtomentum; frontal vitta brownish; area between gena and parafacial brownish to orange; scape, pedicel and postpedicel dark brown; palpus orangish. Thorax black in ground color; postpronotum silvery grey microtomentose; prescutum with two broad silvery grey microtomentose vittae; anterior 3/5 of scutum black and posterior 2/5 silvery grey microtomentose; scutellum black; anepisternum and katepisternum silvery grey microtomentose, remaining pleura black. Legs light brown shading to dark brown tarsi. Upper and lower calypters whitish. Wing hyaline; tegula and basicosta brown; veins yellow. Halter whitish. Abdomen brown in ground color with two broad silver bands; syntergite 1 + 2 with brown mid-dorsal depression and silver lateral and ventral parts; tergites 3-4 each with silver band on anterior half, almost divided by brown triangle in the middle; terminalia brown.
Head ( Figure 8A,B,D-F). Frons at narrowest point much narrower than postpedicel, not more than twice as wide as anterior ocellus, 0.072 of head width and 0.15× as wide as compound eye in dorsal view. Outer vertical seta not differentiated. Ocellar setae well developed, proclinate.
Trichia on arista equally long on upper and lower surfaces; longest trichia longer than width of frons at narrowest point. Head ground color entirely blackish, subparafacial area (i.e., portion between genal grove, genal dilation and vibrissal triangle) and frontal vitta area velvety black (not paler than rest of head). Abdominal tergites 3 and 4 with dark color extending forwards as elongate blackish brown triangle on each tergite with apex at anterior margin (separating pale microtomentose basal parts of tergites into two). Dorsal setulae of abdominal tergites 3 and 4 very long, fine and erect. Frons at narrowest point much narrower than postpedicel, not more than twice as wide as anterior ocellus (0.055× head width).

Phyto paratachinoides Crosskey, 1977
-Trichia on arista not as long on lower surface as upper surface and longest trichia shorter than width of frons at narrowest point. Head not entirely black; subparafacial area and frontal vitta orange-red, brick-red or reddish brown (i.e., paler than rest of head). Abdominal tergites 3 and 4 with dark areas slightly triangular but not extending forwards in mid line to anterior margins of tergites (bases of tergites therefore with uninterrupted broad pale microtomentose fasciae). Dorsal setulae of abdominal tergites 3 and 4 short and recumbent or virtually so. Frons at narrowest point only slightly narrower than postpedicel, conspicuously more than twice as wide as anterior ocellus (0.065× head width).

Discussion and Conclusions
Aporeomyia is an Oriental genus characterized by a tripartite postpedicel in males. In our analyses, Aporeomyia elaphocera sp. nov. clusters with Aporeomyia antennalis Pape & Shima, and the genus is retrieved as sister to Kinabalumyia pinax Cerretti & Pape within an Australasian/Oriental clade. Despite the fact that no female specimens of either species have been found to date, the peculiar antennal morphology likely represents a sexually dimorphic character [20]. Moreover, the discovery of a second species of Aporeomyia has given the opportunity to define this genus based on cladistic arguments as detailed above, and the marked difference in postpedicellar morphology between Aporeomyia and Kinabalumyia is considered here as further support for generic separation.
Baniassa is a Saharo-Arabian/Oriental genus, formerly known from only three species restricted in distribution to the Middle East, all of which shared a petiolate wing cell r 4+5 . Baniassa pennata sp. nov. has the wing cell r 4+5 open, thus presenting the plesiomorphic condition for this character; moreover, this new species remarkably broadens the genus concept and expands the distributional range to include the Oriental Region [34]. However, we are fully aware that the monophyly of Baniassa relies on four weak local apomorphies and that molecular data are needed to better circumscribe the genus.
With 26 described species, the genus Phyto, as it is currently defined, is the second most species-rich genus within Rhinophoridae: only Stevenia Robineau-Desvoidy comprises more species (28) [11]. Phyto mostly shows a west-Palaearctic distribution with only four species in the Afrotropical Region, all endemic: Phyto mambilla sp. nov., P. parafacialis Crosskey, P. paratachinoides Crosskey and P. tachinoides (Curran). According to our phylogeny, the phylogenetic position of Phyto mambilla sp. nov. renders Phyto paraphyletic with regard to Baniassa. Despite this result, we opted for a conservative classification, avoiding both the erection of a new genus to accommodate Phyto mambilla sp. nov. as well as the lumping of Phyto, Baniassa and Parazamimus into a single genus, because we consider such action premature, awaiting the imminent results from a comprehensive phylogeny of the Rhinophoridae based on molecular data (Gisondi et al., in prep.). Except for the remarkably different coloration pattern, characterized by white reflecting microtomentum contrasting with the dark integument, the Afrotropical species of Phyto are very similar to their Palaearctic congeners. Moreover, the two included Palaearctic species of Phyto provide insufficient coverage of the morphological diversity of the genus, and an improved taxon sampling is needed to resolve relationships in this group.
Funding: This research received no external funding.