(C) 2012 Svante Martinsson. This is an open access article distributed under the terms of the Creative Commons Attribution License 3.0 (CC-BY), which permits unrestricted use, distribution, and reproduction in any medium, provided the original author and source are credited.
For reference, use of the paginated PDF or printed version of this article is recommended.
The genus Katatopygia gen. n. is proposed for the Boletina erythropyga/punctus-group that was first introduced by
New genus, taxonomy, Gnoristinae, new synonymy, revision, identification key, phylogeny
In two papers
Accordingly, a new genus Katatopygia gen. n. is here proposed and described for the Boletina erythropyga/punctus-group.
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.
Species | Region | ORI | EN | WN | EP | WP |
---|---|---|---|---|---|---|
# species | 1 | 1 | 4 | 4 | 2 | |
Katatopygia antica (Garrett, 1924), comb. n. | – | – | • | – | – | |
Katatopygia antoma (Garrett, 1924), comb. n. | – | – | • | – | – | |
Katatopygia erythropyga (Holmgren, 1883), comb. n. | – | • | – | • | • | |
Katatopygia hissarica (Zaitzev & Polevoi, 2002), comb. n. | – | – | – | • | – | |
Katatopygia laticauda (Saigusa, 1968), comb. n. | • | – | – | – | – | |
Katatopygia magna (Garrett, 1925), comb. n. | – | – | • | – | – | |
Katatopygia neoerythropyga (Zaitzev & Polevoi, 2002), comb. n. | – | – | – | • | – | |
Katatopygia sahlbergi (Lundström, 1906), comb. n. | – | – | • | • | • |
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:
Type material with original labels of species of Katatopygia gen. n. described by
AMNH American Museum of Natural History, New York, USA
CNC Canadian National Museum, Ottawa, Canada
CUIC Cornell University, Ithaca, New York, USA
KMNH Kyushu University Museum, Fukuoka, Japan
MZLU Museum of Zoology, Lund University, Lund, Sweden
NHRS Swedish Museum of Natural History, Stockholm, Sweden
ZMUN Zoological Museum, University of Oslo, Oslo, Norway
All specimens examined were recorded with unique identification codes prefixed by “JKJ–SPM–” in a BIOTA 2.04 database (
Morphological terminology mainly follows
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
A data matrix (Table 2) for phylogenetic reconstruction was constructed using WINCLADA v1.00.08 (
Observed states of morphological characters used in the phylogenetic studies of Katatopygia gen. n., Katatopygia hissarica, Katatopygia laticauda and Katatopygia neoerythropyga are coded based on original descriptions.
Characters | |||||||||||||||||||||||
---|---|---|---|---|---|---|---|---|---|---|---|---|---|---|---|---|---|---|---|---|---|---|---|
Taxon | 1 | 2 | 3 | 4 | 5 | 6 | 7 | 8 | 9 | 10 | 11 | 12 | 13 | 14 | 15 | 16 | 17 | 18 | 19 | 20 | 21 | 22 | 23 |
Docosia sp. A | 2 | 0 | 1 | 2 | 1 | 0 | 0 | 1 | 0 | 0 | 2 | 0 | 0 | 0 | 2 | 0 | 0 | 0 | 0 | 0 | 2 | 0 | 1 |
Coelosia gracilis | 0 | 0 | 1 | 0 | 1 | 0 | 0 | 0 | 0 | 0 | 1 | 0 | 0 | 0 | 0 | 0 | 0 | 0 | 0 | 2 | 0 | 0 | 1 |
Gnoriste longirostris | 1 | 1 | 1 | 0 | 0 | 0 | 1 | 1 | 0 | 0 | 2 | 0 | 2 | 0 | 0 | 0 | 0 | 1 | ? | ? | 0 | 0 | 1 |
Boletina gripha | 0 | 0 | 1 | 0 | 0 | 0 | 1 | 0 | 0 | 1 | 1 | 0 | 0 | 0 | 0 | 0 | 0 | 0 | 0 | 0 | 2 | 0 | 1 |
Boletina hedstroemi | 0 | 1 | 1 | 0 | 1 | 0 | 1 | 0 | 0 | 1 | 0 | 0 | 0 | 0 | 2 | 0 | 0 | 0 | ? | ? | 2 | 0 | 0 |
Boletina sciarina | 0 | 0 | 1 | 0 | 0 | 0 | 1 | 0 | 0 | 1 | 0 | 0 | 0 | 0 | 2 | 0 | 0 | 0 | ? | ? | 2 | 0 | 0 |
Boletina trivittata | 1 | 1 | 0 | 0 | 0 | 0 | 0 | 0 | 0 | 1 | 2 | 0 | 0 | 0 | 2 | 0 | 0 | 0 | 0 | 2 | 1 | 0 | 0 |
Katatopygia antica | 0 | 1 | 0 | 0 | 0 | 1 | 0 | 1 | 1 | 0 | 1 | 2 | 1 | 1 | 0 | 1 | 1 | 1 | 1 | 1 | 0 | 1 | 0 |
Katatopygia erythropyga | 0 | 1 | 1 | 1 | 0 | 1 | 0 | 1 | 1 | 0 | 1 | 0 | 1 | 1 | 1 | 1 | 0 | 1 | 1 | 1 | 0 | 0 | 0 |
Katatopygia antoma | 0 | 1 | 0 | 0 | 0 | 1 | 0 | 1 | 1 | 0 | 1 | 2 | 1 | 1 | 0 | 1 | 1 | 0 | 1 | 1 | 0 | 1 | 0 |
Katatopygia sahlbergi | 0 | 0, 1 | 0 | 0 | 0 | 1 | 0 | 1 | 1 | 0 | 1 | 1 | 1 | 1 | 0 | 1 | 1 | 1 | 1 | 1 | 0 | 1 | 0 |
Katatopygia laticauda | ? | 0 | 0 | 0 | 0 | 1 | ? | ? | 1 | 0 | 1 | 1 | 1 | 1 | 0 | 1 | 1 | 1 | ? | ? | 0 | 0 | 0 |
Katatopygia hissarica | ? | 1 | 1 | 1 | ? | 1 | ? | ? | 1 | 0 | ? | 0 | 1 | 1 | 1 | 1 | ? | 1 | ? | ? | 0 | 0 | 0 |
Katatopygia neoerythropyga | ? | 1 | 1 | 0 | ? | 0 | ? | ? | 1 | 0 | 1 | 2 | 1 | 0 | 1 | 1 | ? | ? | ? | ? | 0 | ? | 0 |
Katatopygia magna | 0 | 0 | 1 | 1 | 0 | 1 | 0 | 1 | 1 | 0 | 1 | 0 | 1 | 1 | 1 | 1 | 0 | 1 | 1 | 1 | 0 | 0 | 0 |
1. Thorax with: dorsocentrals present = 0; dorsocentrals absent = 1; dispersed setae = 2.
2. Mesonotal stripes: indistinct or absent = 0; distinct = 1.
3. Costa: ending at R5 termination = 0; produced beyond R5 = 1.
4. Sc: non setose = 0; with a few apical setae = 1; mostly setose = 2.
5. CuA-stem: without setae = 0; with setae = 1.
6. Pale abdominal markings: absent = 0; present = 1.
7. Medial fold line of abdominal sternites: absent = 0; present = 1.
8. Tergite VIII: not bearing setae = 0; bearing setae = 1.
9. Male terminalia: not dorsoventrally flattened = 0; dorsoventrally flattened = 1
10. Tergite IX: small, not covering most of gonocoxites and gonostylus = 0; large, covering most of gonocoxites and gonostylus = 1.
11. Gonocoxites: separated = 0; ventrally connected, but not fused = 1; ventrally fused = 2.
12. Gonocoxites: not projected mesocaudally = 0; moderately projected mesocaudally = 1; strongly projected mesocaudally = 2.
13. Hypandrial lobe: vestigial or absent = 0; weakly sclerotized = 1; heavily sclerotized = 2.
14. Gonostylus: without apical processus = 0; with apical processus = 1.
15. Gonostylus: without strong setae on interior surface = 0; with one strong seta on interior surface = 1; with two or more strong setae on interior surface = 2.
16. Apex of gonostylus: without retinacula = 0; with retinacula = 1.
17. Parameres: without microtrichia = 0; with microtrichia = 1.
18. Parameres: paired dorsally = 0; fused into one rod dorsally = 1.
19. Sperm sac: weakly developed or hyaline = 0; well developed and scerotized = 1.
20. Gonocoxal apodeme: vestigial or absent = 0; weakly sclerotized = 1; heavily sclerotized = 2.
21. Cerci: not bearing retinacula = 0; with retinacula evenly distributed = 1; retinacula arranged in lines = 2.
22. Tergite IX: without mesial suture = 0; with mesial suture = 1.
23. Gonostylus: simple = 0; branched = 1.
The data matrix was analysed using parsimony in NONA v2.0 (
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 (
Phylogeny of Katatopygia gen. n. Strict consensus tree (L 53; CI 58; RI 74) calculated from the four most parsimonious trees (L 49; CI 63; RI 79) obtained with the program NONA, with all unambiguous character changes shown and unsupported nodes collapsed. Numbers above branches indicate Jackknife support above 50%. Numbers above hatch marks (black = unique, open = homoplasious) refer to characters, numbers under hatch marks refer to a state changes to the state indicated.
The monophyly of Katatopygia is supported by two unique and one non-unique synapomorphies (character and states given in parenthesis), viz.: 1) Male terminalia dorsoventrally flattened (#9:1), 2) gonostylus with apex covered with retinacula (#16:1), and 3) gonostylus simple (#23:0).
The genus Katatopygia has the parameres fused into one dorsal rod (#18:1) in all species except Katatopygia antoma where paired parameres are retained. This is interpreted as a secondary reversal and this character is here a synapomorphy shared with Gnoriste.
Among the Katatopygia species Katatopygia neoerythropyga is found as the sister-group to the other species. This clade is supported by two synapomorphies viz.: 1) pale abdominal markings present (#6:1) and 2) gonostylus with an apical processus (#14:1). This clade is further subdivided into two distinct clades. One includes the ‘erythropyga-like’ species (Katatopygia erythropyga, Katatopygia hissarica and Katatopygia magna)that is moderately(58 %) supported by Jackknifing and has one synapomorphy; Sc with a few apical setae (#4:1). The other clade includes the ‘salhbergi-like’ species (Katatopygia sahlbergi, Katatopygia antica, Katatopygia antoma, and Katatopygia laticauda)that is moderately(65%) supported by Jackknifing and has three synapomorphies, viz.: 1) costa ending at R5 termination (#3:0, a character state also found in Boletina trivittata), 2) gonocoxites moderately projected mesocaudally (#12:1), and 3) parameres covered with microtrichia (#17:1).
The erythropyga-clade is unresolved, whereas the sahlbergi-clade is fully resolved with Katatopygia laticauda being the sister-group to the remaining species that are united by having tergite IX with a mesal suture (#22:1). Katatopygia antica and Katatopygia antoma share one synapomorphy; gonocoxites strongly projected mesocaudally (#12:2, this state is also found in Katatopygia neoerythropyga).
The data matrix and trees are deposited in the Dryad Data Repository at https://doi.org/10.5061/dryad.682t7442
Systematicsurn:lsid:zoobank.org:act:1A68AEFE-FE7E-4D92-BC9B-430B52D6979A
Boletina sahlbergi Lundström, 1906: 14(type deposited in MZHF)
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, bearing an apical processus (except in Katatopygia neoerythropyga); parameres fused dorsally into one caudally directed rod (with one exception, Katatopygia 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.
Habitus photos of species of Katatopygia gen. n. Lateral view A Katatopygia erythropyga (Holmgren, 1883), male B Katatopygia erythropyga (Holmgren, 1883), female. C. Katatopygia antoma (Garrettt, 1924), male D Katatopygia antoma (Garrettt, 1924), female E Katatopygia sahlbergi (Lundström, 1906), male F Katatopygia sahlbergi (Lundström, 1906), female
Comparison of Katatopygia gen. n. and Boletina Staeger s. str.
Character | Katatopygia | Boletina s. str. |
---|---|---|
Laterotergite | bare | bare or setose |
Median fold line on abdominal sternites | absent | present or absent |
Male terminalia | broad and dorsoventrally flattened | shaped differently |
Male parameres | fused (one exception Katatopygia antoma) | paired |
Male cercus | large, not bearing retinacula | smaller, bearing retinacula |
Male tergite IX | small | large |
Male gonostylus | with apical processus (one exception Katatopygia neoerythropyga) | without apical processus |
Apex of male gonostylus | bearing retinacula | not bearing retinacula |
Female cercus | one segmented | two segmented (except in Boletina abdita and Boletina oviducta) |
Adults: Medium sized, slender with long abdomen, body length 4.5–6.5 mm (Fig. 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.
Morphology of Katatopygia gen. n. [Katatopygia sahlbergi (Lundström, 1906)] A Head, frontal view. B Thorax, lateral view C Front leg D Apex of antenna E Base of antenna. Abbreviations: anepist = anepisternum; anepm = anepimeron; aprnt= anteprononun; ap spur = apical spur; clyp = clypeus; cx 1 = forecoxa; cx 2 = midcoxa; cx 3 = hindcoxa; eye = compound eye; fc = face; fl =flagellar segment; fr fur = frontal furrow; fr tub = frontal tubercle; l oc = lateral ocelus; l cerv scl= lateral cervical sclerite ; ltg = laterotergite; m oc = medial ocellus; mtg = mediotergite; ped = pedicel; plp =palpomere; proepm = proepimeron; sc = scutum; sctl = scutellum; tars 1 = tarsomere one; tb = tibia.
Wings (Fig. 5A–B). Wing membrane unspotted, yellow tinged with dense, irregular arranged microtrichia and no macrotrichia. Crossvein h bare; costa, R1, and R5 with both dorsal and ventral setae; M1, M2, CuA1 and CuA2 with dorsal setae; subcosta bare or with a few setae on distal part; ta, tb, M-petiole, CuA-petiole, A1 and A2 without setae; C ending in, or slightly produced beyond apex of R5; Sc ending in C before or in level with base of Rs; Sc2 present, but may be reduced; R4 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; A1 ending at or slightly before CuA-fork; A2 indistinct and short.
Wing photos of Katatopygia gen. n. A Katatopygia sahlbergi (Lundström, 1906) B Katatopygia antoma (Garrettt, 1924)] Abbreviations: A1 = anterior anal vein; A2 = posterior anal vein; C = Costal vein; CuA1 and CuA 2 = anterior branch of cubitus; CuA–pet = petiole of CuA; h = humeral; M–pet = petiole of media; M1 and M2 = branches of media; R1 = anterior branch of radius; R5 = posterior branch of radius; sc = subcosta; ta = anterior transversal (= crossvein rm); tb = basal transversal.
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.
Morphology of male terminalia of Katatopygia gen. n. A Male terminaliaof Katatopygia erythropyga (Holmgren, 1883), dorsal view, T9 removed B Aedagus and parameres of Katatopygia erythropyga (Holmgren, 1883), lateral view. Abbreviations: aed = aedeagus; aed ap= aedeagal apodeme; ap pro = apical processus of gonostylus; ej ap =ejaculatory apodeme; gc II = section II of gonocoxite; gc III = section III of gonocoxite; gc ap = gonocoxal apodeme; gst = gonostylus; hyp lb = hypandrial lobe; par = paramere; par ap = parameral apodeme; vas def = vas deferens.
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.
Female terminalia of Katatopygia gen. n. A, B Katatopygia erythropyga (Holmgren, 1883) C, D Katatopygia antoma (Garrett, 1924) E, F Katatopygia sahlbergi (Lundström, 1906) [A, C and E in lateral view, B, D and F in ventral view]. Abbreviations: cerc = cereus; gc =gonocoxite; gp =gonapophysis; st = sternite; tg = tergite.
Larvae unknown.
The Nordic species are most abundant in boreal Taiga and subarctic environments, and are possibly strictly boreal-montane. The adults, at least of Katatopygia sahlbergi, seem to be attracted by light, which could suggest nocturnal activity. Larval habitats are unknown for all species in the genus.
A mainly Holarctic genus with the exception of Katatopygia laticauda (Saigusa, 1968), comb. n. described from Taiwan in the Oriental Region (
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.
The species of Katatopygiahttp://sciaroidea.info/taxonomy/41721
Fig. 1A, 8A–CMost similar to Katatopygia antoma, with which it shares the projected dorsomesal corners of the male gonocoxites. Distinguished from Katatopygia antoma by having parameres fused, a small median tooth on hypandrial lobe and brown tip of halter.
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.
Thorax brown with distinct, dark brown mesonotal stripes, humeral area yellow. Antepronotum brown; anepisternum brown; preepisternum brown; laterotergite brown; mediotergite brown. Halter pale with apical part of knob brown.
Wings weakly brownish tinged; veins yellowish brown; stem of M approximately 1.7 times the length of ta; Sc2 present; Sc bare and ending in C at or slightly before base of Rs; C ending at apex of R5.
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.
Female. Coloration as in male, with brown tip of halter. Terminalia not studied.
Katatopygia antica (Garrett, 1924) A Male terminalia, ventral view B Hypandrial lobe C Gonostylus, dorsal view.
Nearctic: Canada, British Columbia.
Only known with the type material.
Syntype series. Canada: B. C. Michel, Wilson Creek. 21 Sep (year unknown pre 1925), leg. C. Garrett – 2 males (CNC, 1 pinned, JKJ-SPM-057739, and 1 pinned with abdomen mounted on separate slide, JKJ-SPM-057740); 24 Sep (year unknown pre 1925), leg. C. Garrett – 3 females (CNC, pinned, JKJ-SPM-057743-45).
http://sciaroidea.info/taxonomy/41722
Figs 1C, 3C–D, 5B, 7C–D, 9A–GMost similar to Katatopygia antica, with which it shares the projected dorsomesal corners of the male gonocoxites. Distinguished from Katatopygia antica by having two parameres, hypandrial lobe without median tooth and pale halter.
Male. Wing length 4.5 mm.
Head brown; palps and mouthparts pale. Antenna with scape, pedicel and basal part of first flagellomere pale, rest of flagellum brown.
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; Sc2 present; Sc bare and ending in C slightly before base of Rs. C ending in apex of R5.
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.
Katatopygia antoma (Garrettt, 1924). A Male terminalia, ventral view B Male terminalia, dorsal view with tergite 9 removed C Male terminalia, caudal view D Aedeagus and parameres, lateral view E Apex of gonostylus, dorsal view F. Hypandrial lobe G Aedeagus and parameres dorsal view.
Nearctic, known from Canada, British Columbia and USA, Alaska.
Syntype series. Canada: BC, Cranbrook, 24 Sep 1922, leg. C. Garrett – 1 female (CNC, pinned, JKJ-SPM-057751); Michel, Wilson Creek. 1 Sep (year unknown pre 1925), leg. C. Garrett – 1 female (CNC, pinned, JKJ-SPM-057753); 24 Sep (year unknown pre 1925), leg. C. Garrett – 3 males (CNC, 1 pinned with cleared terminalia in glycerine, JKJ- SPM-057747, 2 pinned, JKJ- SPM-057748-49), 4 females (CNC, pinned, JKJ-SPM-057755&59-61); 27 Sep (year unknown pre 1925), leg. C. Garrett – 1 female (CNC, pinned, JKJ-SPM-057756); 28 Sep (year unknown pre 1925), leg. C. Garrett – 1 female (CNC, pinned, JKJ-SPM-057757); 2 Oct (year unknown pre 1925), leg. C. Garrett (CNC, pinned, JKJ-SPM-057750); locality and date unknown, marked T.112 – 1 female (CNC, pinned, JKJ-SPM-057752); locality and date unknown, labelled 1815 – 1 female (CNC, pinned, JKJ- SPM-057758).
USA: Alaska, Palmer, 13 Jul 1964 (Leg. K. M. Sommerman) – 2 males, 1 female (MZLU, in alcohol, JKJ-SPM-034388-89).
http://sciaroidea.info/taxonomy/41709
Figs 3A–B, 6A–B, 7A–B, 10A–GMost similar to Katatopygia magna and Katatopygia 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.
Male. Wing length 5.5 mm.
Head blackish brown; mouthparts and palps yellow. Antenna with scape, pedicel and basal part of first flagellomere pale yellow, rest of flagellum brown.
Thorax with three distinct, black mesonotal stripes on yellow ground, humeral area yellow; 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; M-petiole approximately 1.5 times the length of ta; Sc2 present; Sc ending in C slightly before Rs; Sc bearing a few setae on apical portion; C ending beyond apex of R5.
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. 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.
Katatopygia erythropgya (Holmgren, 1883) A Male terminalia, ventral view B Male terminalia, dorsal view C Male terminalia, caudal view D Tergite 9 and cerci, dorsal view E Gonostylys, dorsal view F Aedagus and paramere, dorsal view G Hypandral lobe.
Holarctic, with records from north-western USA (Idaho) (
The species has been confused with Katatopygia sahlbergi and there are some records e.g. from the Alps and Siberia that at least partly refer to the latter species (Zaitzev and Polevoi 2001).
We were not able to locate the holotype of Boletina longicornis in the Johannsen collection at Cornell University (CUIC), nor can it be found in American Museum of Natural History (AMNH) (V. Blagoderov pers. com.) and should probable be regarded as lost.
NORWAY: FV, Alta, Elby, Valsetmoen, sandy slope, 10 Jun-6 Jul 1995 (ZMUN, leg. L.O. Hansen & H. Rinden) – 1 male. SWEDEN: LU, Jokkmokk, Messaure, 2 Sep-4 Oct 1971 (MZLU, leg. K. Möller) – 3 males; Luottåive NR, 28 km S Jokkmokk, 14 Jul-18 Aug 2004 (MZLU, leg. K. Hedmark & J. Kjærandsen) – 1 male; Gällivare, Haapavaara/Annavaara, 8 km WNW Vettasjärvi, 1 Jun-26 Jul 1994 (MZLU, leg. R. Rova) – 1 female, 3 male; SÖ, Haninge, Tyresta National Park, 19 Jun-28 Jul 2000 (NHRS , leg. B. Viklund) – 1 male; TO, Kiruna, Abisko, 150–500 m W Naturv. stn., 18–25 Aug 1975 (MZLU, leg. K. Möller) – 1 male; LF-05, 150–500 m W Naturv. stn., 28 Jun-5 Jul 1976 (MZLU, leg. K. Möller) – 2 males; above tree limit 26 Jun-15 Jul 2006 (NHRS, Leg. Swedish Malaise Trap Project) – 1 female, 10 males; VB, Skellefteå, Stenträsk, Björnhultet Domänreservat, 17 May-17 Oct 1997 (NHRS, leg. B. Viklund) – 4 males.
http://sciaroidea.info/taxonomy/41710
.Very similar to Katatopygia erythropyga, from which it can be separated only on details of the male terminalia. Zaitzev & Polevoi (2002) used four key characters to distinguish them: 1) Apical process of the gonostylus slightly bolder and with more developed unsclerotized area around base of this process; 2) Dentations on the inner surface of gonostylus being restricted to distal part, not reaching the curve basally; 3) Apical part of tergite IX less sclerotized; 4) Details of aedeagus as figured by them.
The species is known only from the holotype from Tadzhikistan.
The species limit between Katatopygia erythropyga and Katatopygia hissarica seems vague, and it is possibly that Katatopygia hissarica will fall inside the variation of Katatopygia erythropyga when a wider range of material is studied.
The holotype is deposited in the Zoological Institute in St. Petersburg, Russia – not studied.
http://sciaroidea.info/taxonomy/41790
Very similar toKatatopygia sahlbergi from which it can be separated by the following characters (from
Taiwan, only known with the holotype.
The holotype is deposited in the Kyushu University Museum, Japan – not studied.
http://sciaroidea.info/taxonomy/41796
Figs 1B, 11A–DKatatopygia magna is most similar to Katatopygia erythropyga and Katatopygia 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.
Male. Wing length 6.0 mm.
Head blackish brown; palps yellow. Antenna with scape brown, pedicel and flagellum yellow.
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; Sc2 present; Sc ending in C clearly before Rs; Sc bearing a few setae on apical portion; C ending beyond apex of R5.
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.
Female unknown.
Katatopygia magna (Garrett, 1925) A Male terminalia, ventral view B Male terminalia, dorsal view, gonostylus omitted C Male terminalia, caudal view, gonostylus omitted D Gonostylus, dorsal view.
Nearctic: Canada, British Columbia.
Known only from the holotype.
Holotype male. Canada: BC, Fernie, 24 Jul (year unknown pre 1925), leg. C. Garrett (CNC, pinned with cleared terminalia in glycerine in microtube on same pin, JKJ-SPM-057738).
http://sciaroidea.info/taxonomy/41711
Most similar to Katatopygia antica and Katatopygia 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 Katatopygia 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.
The species is known only from the Yamal peninsula north in West Siberia.
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).
The holotype is deposited in the A.N. Severtzov Institute of Ecology and Evolution in Moscow, Russia – not studied.
http://sciaroidea.info/taxonomy/41712
Figs 3E–F, 4A–E, 5A, 7E–F, 12A–FMost similar to Katatopygia laticauda from which it can be distinguished by having wing vein Sc2 present and M-pet longer than ta, and in coloration with distinct and separated mesonotal stripes and paler coxae where at most hind coxa are darkened.
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; Sc2 present; Sc ending in C slightly before Rs; C ending at apex of R5.
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.
Female. Body length 6.0-6.5 mm; wing length 6.0–6.5 mm.
Coloration as male.
Terminalia. Tergite IX short with rounded apicolateral corner; sternite VIII and gonocoxite VIII long and narrow, bearing about 6 strong apical setae; gonapophysis IX long and projected into a pointed apex.
Katatopygia sahlbergi (Lundström, 1906) A Male terminalia, ventral view B Male terminalia, dorsal view, tergite 9 removed C Male terminalia, caudal view D Tergite 9 and cerci, dorsal view E Aedeagus and parameres dorsal view F Aedeagus and parameres, lateral view.
Holarctic. Possibly boreal-alpine with records from Scandinavia, northern parts of European Russia, The Alps, West Siberia (Zaitzev & Polevoi 2002), Japan (
The proposed synonymy of Katatopygia sahlbergi and Boletina punctus is based on the study of type material of Boletina punctus and Nordic material of Katatopygia sahlbergi. Katatopygia sahlbergi has been confused with Katatopygia erythropyga and there are some records of the latter species that at least partly refer to Katatopygia sahlbergi (Zaitzev & Polevoi 2002).
NORWAY: FV, Alta, Detsika, Buolamalia, 6 Aug-25 Sep 1996 (ZMUN, leg. L. O. Hansen & H. Rinden) – 2 males; STI, Oppdal, Kongsvoll, 19-26 Jul 1995 (MZLU, leg. J. Skartveit) – 1 male; Kongsvoll, Sprænbekken, 16 Aug-19 Sep 1994 (MZLU, leg. J. Skartveit) – 1 male; SWEDEN: LU, Jokkmokk, Kaltisbäcken 1 km NNE Messaure, 21 Jun-12 Jul 2004 (MZLU, leg. J. Kjærandsen & K. Hedmark) – 1 female, 10 males; 12 Jul-17 Aug 2004 (MZLU, leg. J. Kjærandsen & K. Hedmark) – 3 males; above parking lot, 12 Oct 1997 (MZLU, leg. S. Lundberg) – 1 male; Messaure, 2 Sep-4 Oct 1971 (MZLU, leg. K. Möller) – 14 males; Porsitjärn/Porsi VVO, 1.5 km SE Vuollerim, 6 May-13 Aug 2004 (MZLU, leg. M. Karström) – 2 females, 2 males; 15 Jun-1 Jul 2005 (MZLU, leg. K. Hedmark & M. Karström) – 2 males; 1-16 Jul 2005 (MZLU, leg. K. Hedmark & M. Karström) – 1 male; Tapmokbäckravinen, 12 km SSE Vuollerim, 16 Jun 2004 (MZLU, leg. J. Kjærandsen) – 1 male; Bombmurkleskogen VVO, 4 km SSE Messaure, 17 Jun 2004 (MZLU, leg. J. Kjærandsen) – 1 male; 21 Jun 2004 (MZLU, leg. J. Kjærandsen) – 1 male; 7-19 Jul 2005 (NHRS, leg. Swedish Malaise Trap Project) – 2 males; Luottåive NR, 28 km S Jokkmokk, 14 Jul-18 Aug 2004 (MZLU, leg. K. Hedmark & J. Kjærandsen) – 2 males; 18 Aug-20 Sep 2004 (MZLU, leg. K. Hedmark & J. Kjærandsen) – 4 females, 2 males; Gällivare, Haapavaara/Annavaara, 8 km WNW Vettasjärvi, 1 Jun-26 Jul 1994 (NHRS, leg. R. Rova) – 1 male; Jokkmokk, Bombmurkleskogen VVO, 9-25 Sep 2005 (NHRS, leg. Swedish Malaise Trap Project) – 1 male; 25 Sep-13 Oct 2005 (NHRS, leg. Swedish Malaise Trap Project) – 1 female, 3 males; TO, Kiruna, Abisko, 14-20 Jul 1975 (MZLU, leg. K. Möller) – 4 males; GF-02, 150-500 m W Naturv. stn., 10-25 Jul 1975 (MZLU, leg. K. Möller) – 1 male; LF-01, 150-500 m W Naturv. stn., 6-20 Jun 1975 (MZLU, leg. K. Möller) – 2 males; 22-29 Sep 1975 (MZLU, leg. K. Möller) – 1 male; 29 Sep-6 Oct 1975 (MZLU, leg. K. Möller) – 1 male; LF-02, 150-500 m W Naturv. stn., 21-28 Jul 1975 (MZLU, leg. K. Möller) – 1 male; 28 Jul-4 Aug 1975 (MZLU, leg. K. Möller) – 2 males; 29 Sep-6 Oct 1975 (MZLU, leg. K. Möller) – 1 male; LF-03, 150-500 m W Naturv. stn., 25 Sep-6 Oct 1975 (MZLU, leg. K. Möller) – 13 males; 6-20 Oct 1975 (MZLU, leg. K. Möller) – 2 males; 20-27 Oct 1975 (MZLU, leg. K. Möller) – 1 male; LF-04, 150-500 m W Naturv. stn., 22-29 Sep 1975 (MZLU, leg. K. Möller) – 1 female, 2 males; LF-05, 150-500 m W Naturv. stn., 6-20 Oct 1975 (MZLU, leg. K. Möller) – 1 female, 11 males; 20-27 Oct 1975 (MZLU, leg. K. Möller) – 1 male; LF-06, 150-500 m W Naturv. stn., 4-11 Aug 1975 (MZLU, leg. K. Möller) – 1 male; 1-8 Sep 1975 (MZLU, leg. K. Möller) – 1 male; 15-22 Sep 1975 (MZLU, leg. K. Möller) – 3 males; 22-29 Sep 1975 (MZLU, leg. K. Möller) – 2 females, 11 males; 29 Sep-6 Oct 1975 (MZLU, leg. K. Möller) – 24 males; LF-07, 150-500 m W Naturv. stn., 8-15 Sep 1975 (MZLU, leg. K. Möller) – 1 male; 29 Sep-6 Oct 1975 (MZLU, leg. K. Möller) – 14 males; 6-20 Oct 1975 (MZLU, leg. K. Möller) – 2 males; LF-08, 150-500 m W Naturv. stn., 1-8 Sep 1975 (MZLU, leg. K. Möller) – 1 male; LF-09, 150-500 m W Naturv. stn., 21-28 Jul 1975 (MZLU, leg. K. Möller) – 1 male; 8-15 Sep 1975 (MZLU, leg. K. Möller) – 1 male; 22-29 Sep 1975 (MZLU, leg. K. Möller) – 4 males; 29 Sep-6 Oct 1975 (MZLU, leg. K. Möller) – 14 males; 6-20 Oct 1975 (MZLU, leg. K. Möller) – 1 female, 10 males; LF-10, 150-500 m W Naturv. stn., 22-29 Sep 1975 (MZLU, leg. K. Möller) – 2 males; LSF, 15 Aug-1 Sep 1975 (MZLU, leg. K. Möller) – 1 male; 1-15 Sep 1975 (MZLU, leg. K. Möller) – 2 males; Abisko, Stordalen NR, 9-24 Jul 1975 (MZLU, leg. K. Möller) – 5 males; 7-14 Aug 1975 (MZLU, leg. K. Möller) – 3 males; 4-11 Sep 1975 (MZLU, leg. K. Möller) – 2 males; above tree limit 26 Jun.-15 Jul. 2006 (NHRS, leg. Swedish Malaise Trap Project) – 4 males.
Boletina punctus
Holotype male. Canada: BC, Creston 4 Jul (year unknown pre 1926), leg B. C. D. Garrett (CNC, pinned with terminalia mounted on separate slide, JKJ-SPM-057764). Paratypes. Same data as holotype, marked as allotype – 1 female (CNC, pinned, JKJ- SPM-057778), same data as holotype – 3 females (CNC, pinned, 1 with abdomen in glycerine in microtube on same pin, JKJ- SPM-057779-81), 13 males (CNC, pinned, 2 with terminalia mounted on separate slide, JKJ-SPM-057765-77).
The key is partly based on the key from
1 | Gonostylus with distinct apical processus (cf. Fig. 6A). Abdominal tergites often with pale apical bands | 2 |
– | Gonostylus without distinct apical processus ( |
Katatopygia neoerythropyga |
2 | Costa extending clearly beyond R5-termination. Gonostylus curved inwards, with strong seta on interior surface (cf. Fig. 10E). Sc with a few setae on apical portion. Parameres fused and bare (Fig. 6B) | 3 |
– | Costa ending at or slightly beyond R5-termination (Fig. 5A–B). Gonostylus straighter, without strong seta on interior surface (cf. Fig. 8C). Sc bare. Parameres forked or fused, covered with microtrichia (cf. Figs 9G, 12F) | 5 |
3 | Dentations on the inner surface of gonostylus reaching the curve. Apical part of tergite IX well sclerotized | 4 |
– | Dentations on the inner surface of gonostylus restricted to distal part, not reaching the curve ( |
Katatopygia hissarica |
4 | Gonostylus broadened apically (Fig. 11D). Scutum with dark mesonotal stripes fused | Katatopygia magna |
– | Gonostylus evenly broad (Fig. 10E). Scutum with dark mesonotal stripes distinct and separated | Katatopygia erythropyga |
5 | Mesodorsal corners of gonocoxite distinctly projected caudally (cf. Fig. 9A). Hypandrial lobe only shallowly emarginated (Figs 8B, 9F). Parameres forked or fused | 6 |
– | Mesodorsal corners of gonocoxite not projected (Fig. 12B). Hypandrial lobe deeply divided mesially. Parameres fused into single rod | 7 |
6 | Dorsal part of parameres split into two processes caudally (Figs 9D, G). Hypandrial lobe without a small sharp medial tooth (Fig. 9F). Halter pale (Figs 1C, 3B) | Katatopygia antoma |
– | Dorsal part of parameres fused into one rod caudally. Hypandrial lobe with a small sharp medial tooth (Fig. 8B). Tip of halter brown (Fig. 1A) | Katatopygia antica |
7 | M-pet and ta of approximately the same length. Sc2 absent ( |
Katatopygia laticauda |
– | M-pet longer than ta. Sc2 usually present (Fig. 5A). Scutum pale with dark mesonotal stripes distinct and separated, scutum rarely more uniformly brown. At most hind coxa darkened | Katatopygia sahlbergi |
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;
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 (
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 (
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, Boletina trivittata, may also form a separate clade (
We are very grateful to the curator Dr. Scott E. Brooks at Canadian National Museum in Ottawa (CNC) for arranging the loan of Garrett’s type material. Peter Chandler and an anonymous reviewer provided very helpful comments and improved the English of the manuscript. JK is financially supported by The Swedish Taxonomy Initiative (see