Pteremis Rondani, 1856

Genus Pteremis Rondani, 1856 View in CoL View at ENA

Pteremis Rondani, 1856: 124 View in CoL (gender: masculine). Type species: Borborus nivalis Haliday, 1833 View in CoL [= Pteremis fenestralis ( Fallén, 1820) View in CoL ], original designation.

Pteremis View in CoL : DUDA (1918): 28 (as subgenus of Limosina Macquart, 1835 View in CoL ; diagnosis in key); SPULFR (1924): 375 (as subgenus of Leptocera Olivier, 1813 View in CoL ); RICHARDs (1930): 288 (as subgenus of Leptocera View in CoL ; diagnosis); HACKMAN (1969): 204, 208 (as subgenus of Leptocera View in CoL ; phylogenetic notes, biogeography); PAPP (1984): 87 (as genus; Palaearctic catalog); MARsHALL & RICHARDs (1987): 999 (as subgenus of Leptocera View in CoL ; diagnosis in key, illustr.); RoHÁĆFK (1998):479 (diagnosis in key, illustr.); RoHÁĆFK et al. (2001): 210 (as genus, world catalog).

Pterenis View in CoL (incorrect subsequent spelling): SCHINFR (1864a): 334 (mis- -spelling).

Pterennis View in CoL (incorrect subsequent spelling): SCHINFR (1864b): 49 (catalog, misspelling); SCHINFR (1868): 228 (list of genera), see also O’HARA et al. (2011): 154.

Coprobia Lioy, 1864: 1116 View in CoL (gender:feminine). Type species: Copromyza fenestralis Fallén, 1820 View in CoL , monotypy.

Coprobia View in CoL : RICHARDs (1930): 265 (synonymy).

Stenhammaria Duda, 1918: 28 View in CoL (gender:feminine) (as subgenus of Limosina Macquart, 1835 View in CoL ). Type species: Copromyza fenestralis Fallén, 1820 View in CoL , monotypy.

Stenhammaria View in CoL : RICHARDs (1930): 266 (synonymy).

Paraspelobia Duda, 1938: 96 View in CoL (gender:feminine) (as subgenus of Limosina Macquart, 1835 View in CoL ). Type species: Limosina (Paraspelobia) Vlasovi Duda, 1938 , original designation. New junior subjective synonym.

Paraspelobia View in CoL : HACKMAN (1969):208 (as genus; list, biogeography); Ro- HÁĆFK (1983): 227–232 (as genus, redescription, phylogenetic notes, illustr.); PAPP (1984): 87 (Palaearctic catalog); RoHÁĆFK (1998): 485 (as genus; diagnosis in key, illustr.); RoHÁĆFK et al. (2001): 189 (as genus, World catalog).

Note on gender. Because the gender of the name Pteremis View in CoL was not given in the original description ( RoNDANI 1856: 124) the Art. 30.2.4. of the Code is to be applied saying: ‘If no gender was specified or indicated, the name is to be treated as masculine, except that, if the name ends in -a the gender is feminine, and if it ends in -um, -on, or -u the gender is neuter.’ Formerly, judging on ends of names of species listed in Pteremis View in CoL by PAPP (1984: 87), this generic name was incorrectly treated as feminine, which was followed also in the World Catalog of Sphaeroceridae (RoHÁĆFK et al. 2001) View in CoL . However, based on the above statement of the Code the gender of Pteremis View in CoL is masculine as already recognized by FRFY (1947) when describing Pteremis subapterus Frey, 1947 (a synonym of P. fenestralis View in CoL ) from Finland.

Redescription. (1) pvt present but small to hair-like and convergent to crossed, rarely absent;

(2) occe and occi of medium length; vte, vti (usually longest cephalic seta), oc, posterior ors and vi long and robust;

(3) 4, rarely 3 or 5 ifr, middle pair(s) longer and robust, often cruciate;

(4) 2–9 minute (posterior 2 often longer) ads inside and below ors;

(5) g short, as long as or slightly longer than anterior peristomal setula;

(6) frons with dark stripes between orbits, interfrontalia and frontal triangle almost forming M-shaped mark ( Fig. 101 View Figs 98–103 );

(7) medial carina slightly developed;

(8) 2 hu, but the internal reduced to microseta;

(9) 1 long dc in prescutellar position but dc microsetae in front of it more or less enlarged;

(10) ac microsetae in 6–8 rows on suture, the medial prescutellar ac not or very slightly enlarged;

(11) 2 sc, also laterobasal sc long and robust, sometimes longer than apical sc;

(12) 2 or only 1 stpl but only the posterior strong, the anterior reduced, hair-like or absent;

(13) scutellum relatively large, rounded subtriangular to subcircular, flat to slightly convex;

(14) mid trochanter with only short setae

(15) t 2 chaetotaxy ( Figs 55–57 View Figs 55–61 ) dorsally with proximal and distal pair of strong setae and several short setulae;

(16) t 2 chaetotaxy ventrally variable, always with 1 short av seta below middle ( Figs 23 View Figs 22–27 , 57 View Figs 55–61 , 140 View Figs 138–145 , 153 View Figs 152–157 ) but distally with both va and vpa strong ( Figs 23 View Figs 22–27 , 57 View Figs 55–61 ), or va reduced and vpa strong ( Figs 97 View Figs 91–97 , 116 View Figs 112–118 ) or with only 1 long subterminal seta looking like va ( Figs 140 View Figs 138–145 , 153 View Figs 152–157 ) but probably homologous with vpa shifted more distally;

(17) mt 2 ventrally sometimes with 1 enlarged ventral setula ( Figs 57 View Figs 55–61 , 97 View Figs 91–97 , 154 View Figs 152–157 );

Wing (if present) with

(18) Cs 1 with distinctly ( Fig. 78 View Figs 75–80 ) to slightly ( Fig. 156 View Figs 152–157 ) longer setulae than on rest of C but never as strong as those in Leptocera or Rachispoda Lioy, 1864 species;

(19) C distinctly ( Fig. 16 View Figs 16–21 ) to hardly ( Fig. 156 View Figs 152–157 ) extended beyond apex of R

4+5

;

(20) C-index slightly less to more than 1.0;

(21) R 4+5 slightly sinuate, with apex more ( Figs 16 View Figs 16–21 , 102 View Figs 98–103 ) or less ( Fig. 156 View Figs 152–157 ) upcurved, in brachypterous forms slightly curved ( Figs 17–21 View Figs 16–21 );

(22) dm cell of medium length, with distinct but short processes of M and CuA 1 beyond dm-cu ( Figs 16 View Figs 16–21 , 156 View Figs 152–157 ), in brachypterous forms short, with dm-cu often absent ( Figs 17–21 View Figs 16–21 );

(23) A 1 sinuate, only basal part well developed, distally formed by venal fold ( Fig. 53 View Figs 52–54 );

(24) alula relatively small and narrow, with tapered to acute apex;

(25) abdomen with preabdomen relatively broad, with wide terga; female postabdomen relatively narow ( Figs 34–36 View Figs 33–39 ) to very narrow ( Figs 132, 135–136 View Figs 132–137 , 162, 163 View Figs 162–164 ) and telescopically retractile into preabdomen;

(26) male S5 broad and relatively large, always with posteromedial comb of spines ( Figs 26 View Figs 22–27 , 131 View Figs 125–131 , 157 View Figs 152–157 ) and usually also with a pale-pigmented micropubescent area in front of it ( Figs 26 View Figs 22–27 , 74 View Figs 67–74 , 108 View Figs 104–108 );

(27) male postabdomen relatively small (compared to preabdomen), with S6, S7 and S8 fused and forming strongly asymmetrical synsclerite ( Figs 25, 27 View Figs 22–27 ); near right end of S6, there is an enlarged annular sclerite (= modified 6 th right spiracle, Fig. 27 View Figs 22–27 , rs);

(28) S6+7 with transverse dark-pigmented ledge and 1+1 to 2+3 setae; S7 with irregular posteroventral projection bent inside postabdomen ( Fig. 27 View Figs 22–27 );

(29) male S8 saddle-shaped and almost bare, at most with a pair of small setulae ( Fig. 25 View Figs 22–27 );

(30) epandrium small, uniformly setose, without markedly enlarged setae ( Figs 29 View Figs 28–32 , 126 View Figs 125–131 , 158 View Figs 158–161 );

(31) male cerci large, far produced ventrally, with variously modified (usually flattened) apex); and ventromedially separated by narrow incision ( Figs 28 View Figs 28–32 , 104 View Figs 104–108 , 125 View Figs 125–131 , 159 View Figs 158–161 ), each with 1 long seta and a few shorter setae;

(32) medandrium low (short) and broad, with long lateral arms, each connected with posterior part of gonostylus and posteromedially with narrow, dark-pigmented and ventrally usually split keel ( Figs 28 View Figs 28–32 , 125 View Figs 125–131 );

(33) hypandrium genus-specific, short, with very short and strongly asymmetrical (right directed) anteromedial apodeme ( Figs 29, 31 View Figs 28–32 , hap);

(34) gonostylus ( Figs 28–30 View Figs 28–32 ) of complex bilobed structure, composed of larger anterior (more lateral) lobe and of smaller (shorter and more medial) posterior lobe;

(35) anterior lobe of gonostylus characterized by 3 or 4 strong curved ventral setae, a distinct ventral (posteriorly or posteromedially directed and variously modified) projection and an oblique spinulose posterolateral ledge ( Figs 30 View Figs 28–32 , 85 View Figs 81–87 , 127 View Figs 125–131 , 160 View Figs 158–161 );

(36) posterior lobe of gonostylus with robust terminal spine and posteriorly with subconical process armed by a robust seta on apex ( Figs 30 View Figs 28–32 , 86 View Figs 81–87 , 128 View Figs 125–131 , 160 View Figs 158–161 );

(37) aedeagal complex with phallophore, distiphallus and postgonite subequal in length ( Figs 32 View Figs 28–32 , 87 View Figs 81–87 , 130 View Figs 125–131 , 161 View Figs 158–161 );

(38) phallophore short ( Figs 107 View Figs 104–108 , 130 View Figs 125–131 , 161 View Figs 158–161 ) to very short ( Fig. 50 View Figs 46–51 ), laterally somewhat flattened, ventrally more or less pointed, epiphallus never developed;

(39) distiphallus of simple and relatively uniform construction: weakly sclerotized mainly proximoventrally and dorsally, always with a pair of band-like and pale lateral sclerites and dorsal sclerite subapically forked and bent laterally; middle part of dorsal side of distiphallus finely haired ( Figs 32 View Figs 28–32 , 107 View Figs 104–108 , 161 View Figs 158–161 );

(40) postgonite relatively large, more or less sinuate in lateral view, with tapered distal third to half but with apex always dilated and usually rounded ( Figs 32 View Figs 28–32 , 87 View Figs 81–87 , 107 View Figs 104–108 , 161 View Figs 158–161 ), almost bare;

(41) ejacapodeme present but small to very small ( Figs 32 View Figs 28–32 , 130 View Figs 125–131 , 161 View Figs 158–161 );

(42) female postabdomen narrow to very narrow, gradually tapered ( Figs 34–36 View Figs 33–39 ) or suddenly narrowed and subparallel ( Figs 135–137 View Figs 132–137 ) from preabdomen, always telescopically retractile;

(43) female T7 usually simple ( Fig. 35 View Figs 33–39 ), rarely modified ( Fig. 165 View Figs 165–167 );

(44) female T8 with tripartite pigmentation composed of a pale tongue-shaped medial part and larger darker lateral parts ( Fig. 35 View Figs 33–39 , 135 View Figs 132–137 , 165 View Figs 165–167 ), with a few setae only laterally ( Figs 34, 35 View Figs 33–39 , 135, 136 View Figs 132–137 );

(45) female T10 simply plate-shaped ( Fig. 35 View Figs 33–39 ) to somewhat elongate ( Fig. 135 View Figs 132–137 ), with 1 pair, rarely 2 pairs ( Fig. 165 View Figs 165–167 ) of short medial setae;

(46) female S7 normally simple ( Fig. 36 View Figs 33–39 ), rarely posteromedially emarginate ( Fig. 167 View Figs 165–167 );

(47) female S8 smaller than S7, more or less modified, posteromedially always with a group of setae (all short or 2 long) on elevated sockets ( Figs 38 View Figs 33–39 , 59 View Figs 55–61 , 91 View Figs 91–97 , 112 View Figs 112–118 , 142 View Figs 138–145 , 167 View Figs 165–167 );

(48) female S10 relatively large (about as S8), with anterolateral corners more or less projecting (thus anteromedially emarginate, Figs 39 View Figs 33–39 , 60 View Figs 55–61 , 141 View Figs 138–145 , 167 View Figs 165–167 ), rarely also with elongate medial structure ( Fig. 167 View Figs 165–167 );

(49) female genital chamber largely membranous, with spectacles-shaped sclerite ( Fig. 27 View Figs 22–27 ) represented only by pale-pigmented submembranous rings ( Figs 58 View Figs 55–61 , 113 View Figs 112–118 , 143 View Figs 138–145 );

(50) spermathecae (2+1) subovoid to pyriform, with some surface sculptures (tubercles, spines, rings); terminal parts of ducts long, strongly sclerotized ( Figs 37 View Figs 33–39 , 61 View Figs 55–61 , 115 View Figs 112–118 , 134 View Figs 132–137 ) and sometimes curved ( Fig. 164 View Figs 162–164 ) and those of paired spermathecae connected far from their bodies;

(51) female cerci free, relatively robust, subconical ( Fig. 89 View Figs 88–90 ) or dorsoventrally flattened ( Fig. 35 View Figs 33–39 ), rarely shortened ( Fig. 165 View Figs 165–167 ), each usually with 2 long (apical and dorsopreapical) sinuate setae and a few shorter setae.

Discussion. Because the morphology (particularly of the male and female terminalia) of Pteremis has hitherto not been studied in a sufficient detail its relationships to other genera of Limosininae remains poorly understood. DUDA (1918) classified this group (as subgenera Pteremis and Stenhammaria ) within his informal group ‘Hygrophilae’ comprising otherwise the subgenera Collinella Duda, 1918 (= now genus Rachispoda Lioy, 1864 + genus Leptocera Olivier, 1813 ) and Opacifrons Duda, 1918 (comprising also species now in Pseudocollinella Duda, 1924 ). DUDA (1925, 1938, as tribe Hygrophilae ) also maintained this grouping of subgenera and characterized this assemblage by a strong vpa on t 2 and a distinct (reduced only in Pteremis ) ventral seta on mid basitarsus (called by him ‘Metatarsalborste’). Interestingly, HACKMAN (1969: 204) also considered these subgenera (although also erroneously including Archicollinella Duda, 1925 ) to form a natural group and classified them as subgenera of the restricted genus Leptocera . Subsequently, only RoHÁĆFK (1991), when discussing relationships of Leptocera (including Rachispoda as a subgenus), listed Pteremis together with Paraspelobia , Opacifrons , Pseudocollinella and also Phthitia Enderlein, 1938 as groups with more or less distinct affinities to Leptocera . On the other hand, revisionary studies on Leptocera relatives, viz., Phthitia (see MARsHALL & SMITH 1992), Pseudocollinella (by MARsHALL & SMITH 1993) and Opacifrons (see MARsHALL & LANGsTArr 1998) have not indicated Pteremis as a probable close relative of these genera. OKFLY (1974: 49) described and illustrated the puparium of Pteremis fenestralis – not surprisingly it closely resembles those of the two Leptocera species she also studied (OKFLY 1974: 45) including small, shortly palmate anterior spiracles. Moreover, very similarly formed anterior spiracles are also known in the puparium of Rachispoda species (RoHÁĆFK 1991: Fig. 18 View Figs 16–21 ).

The genus Pteremis (in the extended concept as delimited above) seems to be a monophyletic group supported by the following characters (those considered autapomorphic are marked AA, others also apomorphic A), numbered as above: (16) mid tibia with both vpa and va setae (the latter sometimes secondarily reduced or absent) (AA); (26) male S5 with posteromedial comb of spines and with a pale-pigmented micropubescent area in front of it (A); (31) male cerci large, far produced ventrally, and ventromedially separated by narrow incision (A); (33) hypandrium short, with very short asymmetrical (right directed) anteromedial apodeme (AA); (34) gonostylus of bilobed structure, composed of larger complex anterior lobe and of smaller, simpler, posterior lobe (A); (35) anterior lobe of gonostylus characterized by 3 or 4 strong curved ventral setae, a distinct ventral (posteriorly, posteromedially or posteroventrally directed and variously modified) projection and an oblique spinulose posterolateral ledge (AA); (36) posterior lobe of gonostylus with robust terminal spine and posteriorly with subconical process armed by a robust seta on apex (A); (39) distiphallus of simple and relatively uniform construction but always with a pair of band-like lateral sclerites and dorsal sclerite subapically forked and bent laterally (AA); (40) postgonite sinuate in lateral view, tapered distally but with apex always dilated (A); (47) female S8 more or less modified, posteromedially with a group of setae on elevated sockets (A); (50) spermathecae (2+1) subovoid to pyriform, with terminal parts of ducts long, strongly sclerotized, in paired spermathecae connected far from their bodies (AA).

The presence of both va (directed with axis of tibia) and vpa setae on mid tibia in Pteremis is unique within the Limosininae (see below) irrespective of the fact that va can be reduced and/or vpa shifted distally in aberrant species. The form and structure of hypandrium (33) and the anterior lobe of gonostylus (35), the lateral band-like sclerites of distiphallus (39) and long sclerotized distal parts of spermathecal ducts (50) are also important apomorphies of the genus that separate it clearly from related genera. The armature of male S5, with a posteromedial comb of spines (26) also distinguishes Pteremis from allied genera although a similar comb of spines occurs as homoplasy in several unrelated genera of Limosininae (e.g. Spelobia Spuler, 1924 , Minilimosina Roháček, 1983 ).

The dorsal mid tibial chaetotaxy, general construction of the bilobed gonostylus (34) and shape and armature of its posterior lobe (36), enlarged male cercus and weakly sclerotized distiphallus suggest that Pteremis is related to the genera Phthitia , Pseudocollinella , Rachispoda and Leptocera , as in the orginal ‘Hygrophilae’ of DUDA (1918). This could also be true for the sinuate postgonite, athough its dilated to clubbed apex only occurs in some Pseudocollinella species ( MARsHALL & SMITH 1993). However, the sister-group of Pteremis remains in question. Because in Pteremis a number of distinctly plesiomorphic features is retained (e.g. only single postsutural dc seta, setulae on Cs 1 slightly enlarged; the elongate telescopic female postabdomen with a little reduced and/or modified sterna and terga; unmodified female cerci) it perhaps represents a sister-group to a clade including Leptocera , Rachispoda , Pseudocollinella , Opacifrons and Phthitia although the position of Opacifrons as a member of this group is only likely if its gonostylus is secondarily simplified (simply bilobed only in O. maculifrons group, otherwise unilobate, see MARsHALL & LANGsTArr 1998).

Distribution. The genus is only known from the Holarctic Region.

Species included. Pteremis fenestralis ( Fallén, 1820) (type species), P. apterina sp. nov., P. canaria ( Papp, 1977) , P. ferreus sp. nov., P. kaszabi (Papp, 1973) , P. mongolicus (Papp, 1973) , P. pulliceps sp. nov., P. tenebricus sp. nov., P. vlasovi ( Duda, 1938) comb. nov. (all Palaearctic), P. unicus ( Spuler, 1924) , P. wirthi ( Marshall, 1984) (both Nearctic).