Katatopygia gen. n., a monophyletic branch segregated from Boletina (Diptera, Mycetophilidae)

Abstract The genus Katatopygia gen. n. is proposed for the Boletina erythropyga/punctus-group that was first introduced by Garrett (1924, 1925) and currently comprises eight described species. Molecular studies have strongly indicated that this group forms a monophyletic sister-group to a clade consisting of all other Boletina, Coelosia and Gnoriste, and its monophyly is supported by morphological data as well. The new genus includes the following species: Katatopygia antoma (Garrett, 1924), comb. n., Katatopygia antica (Garrett, 1924), comb. n., Katatopygia erythropyga (Holmgren, 1883), comb. n.,Katatopygia hissarica (Zaitzev & Polevoi, 2002), comb. n., Katatopygia magna (Garrett, 1925), comb. n., Katatopygia laticauda (Saigusa, 1968), comb. n., Katatopygia neoerythropyga (Zaitzev & Polevoi, 2002), comb. n. andKatatopygia sahlbergi (Lundström, 1906), comb. n., all transferred from Boletina. Katatopygia sahlbergi is found to be a senior synonym of Boletina punctus Garrett, 1925, syn. n. A phylogeny based on morphological data and using parsimony analysis yielded four most parsimonious trees where the new genus is retrieved as monophyletic with high support. Katatopygia neoerythropyga is found to be the sister-taxon to all other species that form two clades, one with Katatopygia sahlbergi-like species and one with Katatopygia erythropyga-like species. A key to males of Katatopygia is provided.


introduction
In two papers Garrett (1924Garrett ( , 1925 described four closely allied species of Boletina Staeger from North America that he in the latter paper named the punctus-group and provided a sketchy plate with all four species' male gonostyles aligned. Zaitzev and Polevoi (2002) revised the same group under the name erythropyga-group, but were apparently not aware of and failed to include Garrett's species although their revision was intended to be Holarctic. Although traditionally placed in Boletina, species of this group are morphologically quite distinct in several aspects and well separated from other Boletina as well as from Saigusaia Vockeroth and Aglaomyia Vockeroth, both of which Vockeroth (1980) distinguished from Boletina based on morphological characters. The segregation of Saigusaia has later been supported by phylogenetic studies, both morphological (Søli 1997) and molecular (Martinsson et al. 2011), whereas Aglaomyia may be nested within Boletina (Martinsson et al. 2011). Molecular studies have strongly suggested that Boletina as currently delimited is paraphyletic (Baxter 1999;Martinsson et al. 2011) and given support for the B. erythropyga/punctus-group being the sister-group to a clade consisting of all other Boletina, Aglaomyia, Coelosia Winnertz and Gnoriste Meigen (Martinsson et al. 2011).
Accordingly, a new genus Katatopygia gen. n. is here proposed and described for the B. erythropyga/punctus-group. Holmgren (1883) described the first species as Boletina erythropyga Holmgren, 1883, based on material from Novaya Zemlya in northern Russia. Lundström (1906) added Boletina sahlbergi Lundström, 1906from Finnish Lapland, while Johannsen (1911 added Boletina longicornis Johannsen, 1911 from Idaho in USA. Garrett (1924Garrett ( , 1925 described four additional species based on material from British Columbia in Canada, of which Boletina punctus syn. n. is considered here a junior synonym of Boletina sahlbergi. Saigusa (1968) added Boletina laticauda Saigusa, 1968 from Taiwan. Laštovka and Matile (1988) erroneously synonymized Boletina sahlbergi with Boletina erythropyga, leading Zaitzev (1994) to table 1. List of World species of the genus Katatopygia gen. n. All species are being transferred from Boletina Staeger. Their known distribution in faunal regions and subregions is given to the right. Abbreviations: ORI -Oriental Region EN -Eastern Nearctic subregion WN -Western Nearctic subregion WP -Western Palaearctic subregion EP -Eastern Palaearctic subregion.
---• -Katatopygia sahlbergi (Lundström, 1906), comb. n. - describe a new species representing the former. Zaitzev and Polevoi (2002) resolved these confusing species interpretations by reinstating Boletina sahlbergi and suggesting two new synonyms, and finally added two more species. With these changes a group that currently includes eight species can be assigned to Katatopygia gen. n. (Table 1), all being transferred from Boletina s.l.

Material and methods
The examined material was gathered from museum collections and surveys, and mainly consists of the type series of the species described by CBD Garrett (Fig. 1) from Canada, some material from Alaska (USA) and European material from the Nordic region. The following collection acronyms for depositories are used in the text: All specimens examined were recorded with unique identification codes prefixed by "JKJ-SPM-" in a BIOTA 2.04 database (Colwell 2007), and the lists of material examined were extracted from this database. For each species and country the localities are sorted hierarchically within provinces, districts, localities and sites, respectively.
Morphological terminology mainly follows Søli (1997), the term "retinacula" is here used for any assemblage of strong, short and blunt macrosetae. The term "apical processus" is adopted from Zaitzev & Polevoi (2002) and is used for a small appendage that articulates to an unsclerotized area apically on the gonostylus. Terminology of sensillae follows Seifert (1975).
Terminalia were macerated in heated KOH (90°C) and transferred to acetic acid for neutralisation, then to alcohol and finally to glycerine. Most terminalia are preserved in glycerine in micro-vials together with the rest of the specimen, while some specimens are permanently mounted in Canada balsam on slides as outlined by Kjaerandsen (2006). In order to produce plates the terminalia were either photographed in glycerine with a Nikon Digital Sight DS-M5 microscope camera mounted on a Nikon SMZ1500 stereomicroscope, or placed in alcohol gel under a coverslip and photographed through a Nikon Eclipse 50i compound microscope. Series of z-stack photos were taken and combined for extended focus using HELICON FOCUS. The images were digitally edited in ADOBE PHOTOSHOP and GIMP. Scanned sketches, drawn using a drawing tube attached to a Nikon Eclipse 50i compound microscope, were used as templates to produce digital illustration with GIMP.

Phylogenetic analysis
A data matrix (Table 2) for phylogenetic reconstruction was constructed using WIN-CLADA v1.00.08 (Nixon 2002). Characters dealing with structures of the thoracic sclerites, wings, abdominal sclerites and male terminalia were used, with a focus on characters of the terminalia. The characters were either coded as binary (15) or multistate (8). Missing data was coded as "?". All species of Katatopygia were included as the ingroup. Character states for K. laticauda, K. neoerythropyga (Zaitzev & Polevoi, 2002) and K. hissarica (Zaitzev & Polevoi, 2002) were derived from the original descriptions (Saigusa 1968;Zaitzev and Polevoi 2002). As outgroups we used Coelosia gracilis Johannsen, Gnoriste longirostris Siebke, an undescribed species of Docosia cf. gilvipes (Kjaerandsen & Hedmark in prep.)  The following 23 characters were used in the analysis; observed character states are given in Table 2. 1.
The heuristic search produced four most parsimonious trees (L 49; CI 63; RI 79). A strict consensus tree was calculated and is shown in Fig. 2 with all unambiguous character changes and unsupported nodes collapsed. The new genus Katatopygia forms a monophyletic group that is statistically supported (92%) by Jackknifing. This is in accordance with previous molecular studies (Martinsson et al. 2011) although we here end up with a different arrangement among outgroup genera and clades that gain little or no statistical support by the Jackknife analysis.
The genus Katatopygia has the parameres fused into one dorsal rod (#18:1) in all species except K. antoma where paired parameres are retained. This is interpreted as a secondary reversal and this character is here a synapomorphy shared with Gnoriste.
The erythropyga-clade is unresolved, whereas the sahlbergi-clade is fully resolved with K. laticauda being the sister-group to the remaining species that are united by having tergite IX with a mesal suture (#22:1). K. antica and K. antoma share one synapomorphy; gonocoxites strongly projected mesocaudally (#12:2, this state is also found in K. neoerythropyga).
The data matrix and trees are deposited in the Dryad Data Repository at doi: 10.5061/dryad.682t7442 systematics Genus Katatopygia gen. n. urn:lsid:zoobank.org:act:1A68AEFE-FE7E-4D92-BC9B-430B52D6979A http://species-id.net/wiki/Katatopygia http://sciaroidea.info/taxonomy/41708 Type species. Boletina sahlbergi Lundström, 1906: 14(type deposited in MZHF) Diagnosis. The genus consists of medium sized slender Gnoristinae with long abdomen where the males have a very characteristically flattened and dilated terminalia (eg. Fig. 3E). They can be recognized on a combination of the following characters: mouthparts not prolonged; scutum with setae arranged in acrostichals, dorsocentrals and laterals; laterotergite bare; wing with Sc ending in C; M-petiole as long as or longer than ta; CuA-furcation before level of M-fork, approximately level with base of Rs; abdominal sternites with median fold-line absent; male terminalia broad and dorsoventrally flattened, often rotated about 180°; gonostylus large and simple, bear-ing an apical processus (except in K. neoerythropyga); parameres fused dorsally into one caudally directed rod (with one exception, K. antoma, that has pared parameres); cerci large and without retinacula, covered with long trichia; hypoproct well developed; female terminalia with unsegmented cerci. The best characters to separate between Katatopygia and Boletina s.s. are further listed in Table 3.
Head (Fig. 4A). Vertex with scattered setae. Ocelli three, almost in line, the median slightly smaller than laterals, lateral ocelli separated from eye by approximately 1.5 times its diameter; below the ocelli, protuberances present and well sclerotized. Eyes with shallow emargination above antennal base. Frons without setae but with small microtrichia and on lateral parts some stronger microtrichia; frontal furrow well developed and reaching apex of frontal tubercle. Antenna with 14 flagellomers; scape and pedicel with a few scattered setae and short microtrichia (Fig. 4E); flagellomeres long rectangular, densely covered with medium sized setae; apical flagellomere with a somewhat stronger terminal seta (Fig. 4D). Face with scattered setae. Mouthparts not prolonged; clypeus oval to subtriangular and well separated from face, sclerotized and bearing setae; palps with five palpomeres, the first being reduced and easily overlooked, third palpomere with sensillae on inner surface. Thorax (Fig. 4B). Antepronotum fused with proepisternum, bearing some setae, the suture between the sclerites weak. Scutum with setae arranged in acrostichals, dorsocentrals and laterals, otherwise bare. Scutellum with one pair of bristles and scattered setae. Anepisternum, anepimeron latero-and mediotergite all bare.
Wings (Fig. 5A-B). Wing membrane unspotted, yellow tinged with dense, irregular arranged microtrichia and no macrotrichia. Crossvein h bare; costa, R 1 , and R 5 with both dorsal and ventral setae; M 1 , M 2 , CuA 1 and CuA 2 with dorsal setae; subcosta bare or with a few setae on distal part; ta, tb, M-petiole, CuA-petiole, A 1 and A 2 without setae; C ending in, or slightly produced beyond apex of R 5 ; Sc ending in C before or in level with base of Rs; Sc 2 present, but may be reduced; R 4 absent; M-petiole between 1 and 2 times as long as ta; CuA-fork starts proximally of M-fork, approximately at the level of base of Rs; A 1 ending at or slightly before CuA-fork; A 2 indistinct and short. Legs (Fig. 4C). Legs often pale with dark setation; fore and mid coxae with some setae on apical part; trochanter dark; bearing sensillae and a few small setae; femur with numerous setae and no bristles; tibia covered with irregularly arranged setae and with bristles mainly confined to ventral surface; fore tibia with anteroapical depressed area semicircular and densely covered with long microtrichia; apical tibial spur serrated and covered with microtrichia, no apical comb present; tarsus covered with macrotrichia and some stronger setae; claws with a small ventral lobe; empodium pulvilliform.
Abdomen. Pale abdominal markings, when present, situated towards the apices of the tergites. Sternite 1 with a few weak setae apically, all other segments haired; sternites with sublateral fold-lines, median fold-line absent; segment 7 and 8 reduced and retracted into segment 6.
Male terminalia (Fig. 6A-B). Broad and dorsoventrally flattened; often rotated about 180°. Tergite IX rather small and subrectangular, in some species with a mesial sclerotized suture, scattered with setae. Cerci large, rounded to oval, without retinacula, densely covered with long microtrichia. Gonocoxites large, moderately incised ventrally with a hypandrial lobe situated in this incision; hypandrial lobe well developed and more or less branched; gonocoxite bearing scattered macro-and micotrichia, long microtrichia densely covering apical margin. Tergite X present as a weakly sclerotized, short and broad plate situated ventrally, near apex of tergite IX. Hypoproct well developed, situated ventrally to cerci and fused with tergite X, setose and resembling a second segment of cercus. Gonostylus large, unbranched except posessing a tiny apical processus which articulates to a small unsclerotized area and bears 1-2 strong setae, in some species this processus is minute or absent; apex of gonostylus covered with dense retinacula; ventrobasally surface of gonostylus with patch of placoid sensillae; inner surface of gonostylus usually fringed with small dentations. Accessory copulatory appendages joined to gonocoxite through a weakly sclerotized gonocoxal apodeme attached near apex of aedeagus. Aedeagus apically connected with parameres; in most species the parameres are fused dorsally into one caudally directed rod; aedeagus with well developed sperm sacs, to which vas deferens is attached.
Female terminalia (Fig. 7A-F). Tergite VIII well developed, subrectangular. Sternite VIII well developed, entirely fused with gonocoxite VIII that is tapered and bearing several strong setae at apical margin. Tergite IX well developed, shorter than Tergite VIII. Gonapophysis VIII hyaline, indistinct. Gonapophysis IX ventrally divided and retracted into segment VIII, in some species projected into a pointed apex, while in other short and blunt. Tergite X very short, laterally fused with sternite X that is completely divided ventrally and projected caudally. Cerci one-segmented, ovate.
Larvae unknown. Notes on biology. The Nordic species are most abundant in boreal Taiga and subarctic environments, and are possibly strictly boreal-montane. The adults, at least of K. sahlbergi, seem to be attracted by light, which could suggest nocturnal activity. Larval habitats are unknown for all species in the genus.

Distribution.
A mainly Holarctic genus with the exception of Katatopygia laticauda (Saigusa, 1968), comb. n. described from Taiwan in the Oriental Region (Saigusa 1968). The greatest diversity is found in Western North America and in the Eastern Palaearctic with four species in each of the regions (Table 1).
Etymology. Katatopygia is derived from the Greek words katatonis, meaning "broader than high", pygo-, meaning "rump" or "buttock" and the suffix -ia denoting pertaining to. The name refers to the characteristic broad and dorsoventrally flattened terminalia shared by all males in the genus. The name is a noun and is feminine.
Re-description. Male. Wing length 5.0-5.5 mm. Head brown; palps and mouthparts pale. Antenna with scape brown, pedicel and basal part of first flagellomere pale, rest of flagellum brown.
Wings weakly brownish tinged; veins yellowish brown; stem of M approximately 1.7 times the length of ta; Sc 2 present; Sc bare and ending in C at or slightly before base of Rs; C ending at apex of R 5 .
Legs pale with joints darker. Abdomen dark brown often with narrow pale apical bands on tergites II-IV. Terminalia brown. Gonocoxite with dorsomesal corner forming a mesocaudally directed horn-like processus, distinctly more projected than the ventromesal corner. Hypandrial lobe well developed and only shallowly emarginated medially with a small sharp medial tooth. One slender paramer, bearing microtrichia. Tergite IX subrectangular, with a sclerotized mesal suture. The apical processus on gonostylus approximately half as long as the diameter of gonostylus and slightly branched with two strong setae. Interior surface of gonostylus without strong setae.
Thorax pale with 3 distinct, dark brown mesonotal stripes on yellow ground, humeral area pale. Mediotergite with a darker central stripe; preepisternum darker ventrally. Halter whitish.
Wings weakly brownish tinged; stem of M approximately 1.9 times the length of ta; Sc 2 present; Sc bare and ending in C slightly before base of Rs. C ending in apex of R 5 .
Legs pale brown. Abdomen brown often with narrow pale bands on tergites II-III. Terminalia brown. Gonocoxite with dorsomesal corner forming a mesocaudally directed horn-like processus, distinctly more projected than the ventromesal corner. Hypandrial lobe well developed and only shallowly emarginated medially, without medial tooth. Two slender parameres, bearing microtrichia. Tergite IX subrectangular, with a sclerotized mesal suture. The apical processus on gonostylus approximately half as long as the diameter of gonostylus and slightly branched with two strong setae. Interior surface of gonostylus without strong setae.
Female. Coloration as in male except pale apical bands on tergite II-V. Terminalia. Tergite VIII broad with rounded apicolaterally margin; sternite VIII and gonocoxite VIII short and broad with about 6 strong apical setae; gonapophysis IX long and projected into a pointed apex.
Distribution. Nearctic, known from Canada, British Columbia and USA, Alaska. Diagnostic characters. Most similar to K. magna and K. hissarica, but can be distinguished by having distinct and separated mesonotal stripes and on the evenly broad male gonostylus on which the inner dentations are reaching the basal curve.
Head blackish brown; mouthparts and palps yellow. Antenna with scape, pedicel and basal part of first flagellomere pale yellow, rest of flagellum brown.
Wing pale with veins yellowish brown; M-petiole approximately 1.5 times the length of ta; Sc 2 present; Sc ending in C slightly before Rs; Sc bearing a few setae on apical portion; C ending beyond apex of R 5 .
Legs pale yellow with joints darker. Abdomen dark brown with yellow apical bands on tergites II-VI. Terminalia brownish. Gonocoxite with mesal corners not projected; gonostylus evenly broad, angled inwards about 40° and bearing one strong seta on interior surface, dentations on interior surface reaching curve. Apical processus approximately as long as the diameter of gonostylus, bearing one apical seta. Hypandrial lobe deeply forked with four lobes. Dorsal fused paramere rod long, slender and without microtrichia.  (Garrett, 1924) A Male terminalia, ventral view B Hypandrial lobe C Gonostylus, dorsal view.
Tergite IX subrectangular, with 4 stronger setae on apical part and without a sclerotized mesial suture.
Female. Body length 6.0 mm; wing length 5.5 mm. Coloration as male.
Terminalia. Tergite IX broad with sharp apicolateral corner; gonocoxite VIII slightly incised ventrally bearing many strong apical setae; gonapophysis IX short and blunt.
Distribution. Taiwan, only known with the holotype. Type material. The holotype is deposited in the Kyushu University Museum, Japan -not studied. (Garrett, 1925), comb. n. http://sciaroidea.info/taxonomy/41796 Figs 1B, 11A-D Garrett, 1925: 5 Diagnostic characters. K. magna is most similar to K. erythropyga and K. hissarica, but can be distinguished by having fused mesonotal stripes and on the apically broadened gonostylus on which the inner dentations are reaching the basal curve.
Thorax with black mesonotal stripes fused on yellow ground, humeral area yellow; antepronotum brown; anepisternum brown; preepisternum dark with a diffuse pale spot; laterotergite brown; mediotergite dark. Halter pale.
Wing pale with veins yellowish brown; M-petiole approximately 1.8 times the length of ta; Sc 2 present; Sc ending in C clearly before Rs; Sc bearing a few setae on apical portion; C ending beyond apex of R 5 .
Legs pale yellow with joints darker. Abdomen dark brown with yellow apical bands on tergite II-IV. Terminalia yellowish. Gonocoxite with mesal corners not projected; gonostylus with broadened apex; gonostylus angled inwards about 65° and bearing one strong seta on interior surface, dentations on interior surface reaching curve. Apical processus approximately as long as the diameter of gonostylus and bearing one subapical seta. Hypandrial lobe deeply forked with four lobes. Dorsal fused paramere rod long and straight, without microtrichia. Tergite IX subrectangular, without sclerotized mesial suture.

Katatopygia neoerythropyga
Boletina neoerythropyga Zaitzev & Polevoi, 2002: 641 (figs 7-8) Diagnostic characters. Most similar to K. antica and K. antoma from which it can be separated on coloration and details of the male terminalia. Zaitzev & Polevoi (2002) used four key characters to distinguish it from K. erythropyga: 1) absence of the apical process of the male gonostylus; 2) longer stem of M-fork; 3) scape of antenna brown; 4) abdomen uniformly brown.
Distribution. The species is known only from the Yamal peninsula north in West Siberia.
Remarks. The absence of the apical process of the male gonostylus is unique among the known species of Katatopygia and may be regarded as a secondary reduction (see discussion of phylogeny).
Type material. The holotype is deposited in the A.N. Severtzov Institute of Ecology and Evolution in Moscow, Russia -not studied.
Re-description. Male. Body length 4.5-6.5 mm; wing length 4.5-6.0 mm. Head blackish brown; mouthparts and palps yellow. Antenna with scape, pedicel, and basal part of first, in some specimens the whole first and basal part of second, flagellomere pale yellow, rest of flagellum brown.
Thorax in most specimens with 3 distinct, black mesonotal stripes on yellow ground, humeral area yellow, a few specimens with mesonotal stripes indistinct and humeral area brownish; antepronotum pale; anepisternum brown; preepisternum pale with ventral half darker, brown; laterotergite brown with anterior part paler; mediotergite pale with a broad dark central stripe. Halter pale.
Wing pale with veins yellowish brown; stem of M approximately 1.5 times the length of ta; Sc 2 present; Sc ending in C slightly before Rs; C ending at apex of R 5 .
Legs pale yellow with joints darker. Abdomen dark brown, usually with yellow apical bands on tergite I-IV. Terminalia often yellow with dark lateral markings, in some specimens brownish and not distinctly paler than rest of abdomen. Gonocoxite with mesal corners slightly projected. Tergite IX subrectangular, with a sclerotized mesal suture. Paramere simple, strong, blunt and covered with microtrichia. Gonostylus straight with apical processus approximately half as long as the diameter of gonostylus.

Discussion
Resolving phylogeny of the extended Gnoristinae clade is way beyond the scope of this study and the quantitative phylogenies that have been presented so far (e.g. Söli 1997;Baxter 1999;Rindal et al. 2009;Martinsson et al. 2011) are partly conflicting and not very convincing when it comes to stable intergeneric relationships. With some 160 species placed in the rather heterogeneous genus Boletina we still think that it will benefit from being split into subsets of putatively natural entities, and that these taxa will shed new light on the phylogeny of the entire group. The process was initiated by Martinsson et al. (2011) who estimated the first molecular phylogenies focused on Boletina and related genera. The decision to erect the new genus Katatopygia rest largely on the supportive results from Martinsson et al. (2011) where the two European species consistently and with support across different genes were found as the basal sistergroup to all the other species of Boletina, Coelosia and Gnoriste included in the analysis. The unequivocal basal position makes it rather unlikely that our splitting will render Boletina s.s. paraphyletic with respect to Katatopygia. The idea to segregate species of the B. erythropyga/punctus-group from Boletina s.l. is, however, not new and grew out of a long-funded distinct impression that these species form a morphological uniform group with highly specialized male terminalia that don't naturally fit together with the remaining Boletina s.s. The additional morphological analysis presented here was designed to test the monophyly of the extended group of eight Katatopygia species and resolve their interrelationships. The characters included in the analysis were thus chosen mainly for resolving relationships among Katatopygia species, not to resolve relationships among Gnoristinae genera. A few outgroup taxa were, based on the available phylogenies (Søli 1997;Rindal et al 2009;Martinsson et al 2011), selected among genera available to us that have been indicated to be closely related to Boletina s.l. (including Katatopygia). Given the rather limited selection of informative characters found and the few outgroup taxa included the present analysis cannot be given the same credit to support the segregation of Katatopygia as did the molecular study (Martinsson et al. 2011). Focused mainly on terminalia morphology the analysis mainly summarizes those characters we found to be diagnostic for the new genus and gives a first clue to inter-relationships among its species. Accordingly the morphological analysis retrieved a monophyletic Katatopygia with high support whereas relationships among the outgroup taxa are not supported and somewhat contrasting those found by Martinsson et al. (2011).
The segregation of Katatopygia rise new questions related to the increasing number of Boletina "look-alike" genera. Are they forming a monophyletic clade together with Boletina s.s. or rather constitute an assemblage of less related plesiomorphous genera? Another recently described genus, Heamesphaerenotus Saigusa from China (Saigusia 2007), show some affinities with Katatopygia in general appearance and they have some possibly apomorphic characters in common (retinacula on male gonostylus and one-segmented female cercus). But like for Katatopygia, Heamesphaerenotus also show a number of unique apomorphies, notably the expanded eave-like mesonotum (Saigusa 2007) not seen in Katatopygia. Unlike species of Boletina s.s. species of Aglaomiya, Heamesphaerenotus, Katatopygia and Saigusaia all have some form of retinacula on the apical part of the gonostylus, and this can also be found among species of Synapha. The females of Saigusaia and Synapha have two-segmented cerci (e.g. Saigusa 1968) whereas all known females of Katatopygia and Heamesphaerenotus have one-segmented cerci. In fact, Saigusaia seems to be much closer Synapha than to Boletina (Søli 1997;Martinsson et al. 2011) and the general outline of the male terminalia among species of Saigusaia (see e.g. Niu et al. 2008) very much resembles that of some Synapha (see e.g. Kallweit and Martens 1995).
Even after the segregation of Katatopygia, Boletina s.s. remain as a large and somewhat heterogeneous genus. It is noteworthy that the type-species of the genus, B. trivittata, may also form a separate clade (Martinsson et al. 2011) including some 10 morphological similar species. Some aberrant species currently included in Boletina, e.g. B. abdita Plassmann, B. anderschi (Stannius) and B. ovata (Garrett), needs further studies to see if they fit within a restricted definition of the genus.