Zwicknia bifrons ( Newman, 1838 ) Murányi & Gamboa & Orci, 2014

Murányi, Dávid, Gamboa, Maribet & Orci, Kirill Márk, 2014, Zwicknia gen. n., a new genus for the Capnia bifrons species group, with descriptions of three new species based on morphology, drumming signals and molecular genetics, and a synopsis of the West Palaearctic and Nearctic genera of Capniidae (Plecoptera), Zootaxa 3812 (1), pp. 1-82 : 40-44

Zwicknia bifrons ( Newman, 1838) comb. n.

( Figs. 1–2 , 7, 10 , 23 , 32 , 56 , 58 , 62–63 , 66 , 72, 74, 76 , 82–85 , 106 , 114 , 120 , 139–143 , 152–156 , 165 , 169–173 , 185 , 190–192 , 196–197 ).

Chloroperla bifrons Newman, 1838 — Newman 1838: 401. (original description of the adult)

Capnia nigra ( Pictet, 1833) —figures in at least Morton 1896, Hynes 1940, Hynes 1941 and Hanson 1946 (enumerated under the general synonymies) refers to Z. bifrons .

Capnia bifrons ( Newman, 1838) — Kimmins 1947: 261. (stat. rev and comb. n.);—figures in at least Brinck 1949, Kimmins 1950a, Kimmins 1950b, Brinck 1952, Hynes 1955a, Hynes 1958, Illies 1955, Khoo 1964, Lillehammer 1965, Lillehammer 1974, Rupprecht 1976 and Lillehammer 1988 (enumerated under the general synonymies) refers to Z. bifrons ;—drumming signals in Rupprecht 1965, Rupprecht 1968, Rupprecht 1976 (enumerated under the general synonymies) and partly in Rupprecht 1982 ( Fig. 2b ) refers to Z. bifrons .

Capnia bifrons ( Newman, 1838) Capnor race sensu Rupprecht 1997 — Rupprecht 1997: 94. (drumming signals); Enting & Rupprecht 2001: 71 (clarification as the nominal C. bifrons ).

Diagnosis. Male epiproct: Ep-scl wide and blunt in dorsal view, tip upcurved in lateral view; ventral membranous section nearly reach the base in lateral view, apical spines distributed only on the membranous part of the apex. Process of male Tg 9: high, perpendicularly elevated, <1½× wider than Ep-scl, rectangular, and with continuously narrow sides caudally. Males usually produce drumming calls in sequences containing 2–3 calls ( Fig. 165 ), but single calls are also produced sporadically. The male call is a monophasic beat-group with duration of 1400–2400 ms and composed of 7–16 beats repeated with a mean inter-beat interval of 140–230 ms. Inter-beat interval gradually increases during each male call ( Figs. 169–173 , 190 ). Inter-call interval increases as a call sequence proceeds, but its mean value is between 1400–6300 ms. All of these drumming character values refer to signals produced at ambient air temperature between 15–21 o C. The male-female drumming duet is a long sequence of male call—female answer alternation ( Fig. 185 ).

Material examined. HUNGARY: Veszprém County, Bakony Mts., Bakonybél, Gerence Stream at Borostyán Spring , N 47°14.596’ E 17°43.835’, 275 m, 24.02.201 1, leg. D. Murányi , K. M. Orci: 1m ( HNHM; used for drawings Figs. 142 , 155 , for molecular studies as 300995, drumming recorded as 2011/ No. 1, Fig. 170 ) ; Pest County, Pilis Mts., Szentendre, Dömörkapu , Bükkös Stream , N 47°41.751’ E 18°59.904’, 260 m, 23.02.201 0, leg. D. Murányi , K. M. Orci: 7m 4f, 1m larva ( HNHM; one male, three male terminalia, one female and the male larva prepared for SEM, two female and parts of a male terminalia prepared on slides, specimens used for drawings and photos Figs. 1–2 , 7, 10 , 23 , 32 , 56 , 58 , 62–63 , 66 , 72, 74, 76 , 82–85 , 106 , 114 , 120 ), 6m 7f ( HNHM; drummings recorded as 2010/ No. 1–3, 6–9), 4m ( BYUC) , 1m 1f ( GVC; drummings recorded as 2010/ No. 4), 1m 1f ( PZC; drummings recorded as 2010/ No. 5) ; 26.02.201 1, leg. K. M. Orci: 2m ( HNHM; used for molecular studies as 300991, 300978, drummings recorded as 2011/ No. 1–2) ; 14.03.2011, leg. K. M. Orci: 3m ( HNHM; used for molecular studies as 300979, 300986, 300984, drummings recorded as 2011/ No. 3–4, Fig. 169 ) ; 27.03.201 1, leg. K. M. Orci: 1m ( HNHM) ; Pest County, Börzsöny Mts., Szokolya-Királyrét, Szén Stream , N 47°53.672’ E 18°58.706’, 255 m, 05.03.201 1, leg. D. Murányi : 4m ( HNHM; used for drawings Figs. 139 , 152 , for molecular studies as 300990, 300977, 301002, 300980, drummings recorded as 2011/ No. 1–2, 4, Fig. 171 ) ; Heves County, Mátra Mts., Gyöngyössolymos, Monostor Stream , N 47°49.918’ E 19°55.801’, 260 m, 07.03.201 1, leg. T. Kovács: 12m 2f ( HNHM; used for drawings Figs. 140 , 153 , for molecular studies as 300950, 300972, 300976, 300953, 300954, 300975, drummings recorded as 2011/ No. 1–5, Fig. 172 ) ; SERBIA: Zlatibor District, Maljen Mts., Brajkovići , stream N of the village, N 44°02.244’ E 19°54.827’, 445 m, 17.03.201 1, leg. T. Kovács, G. Magos, D. Murányi : 2m 1f ( HNHM; one male used for drawings Figs. 143 , 156 ), 9m ( HNHM; used for molecular studies as 300994, drummings recorded as 2011/ No. 1–7 Fig. 173 ) .

Other material—Records based on morphology: AUSTRIA: Vienna State, Wien, Maurbach , 03.03.195 3, leg. E. Pomeisl: 1m 1f, 3 larvae ( WNHM; male terminalia in microvial) ; Lower Austria and Vienna States , Vienna Woods , many brooks, various dates from February to March, leg. W. Graf ( WGC) ; Styria State, Lanzegg , 16.03.200 4, leg. W. Graf: 15m 22f ( WGC) ; FRANCE: Rhône-Alpes Region, Ardèche, Rivere de Celle , 20.03.195 0, leg. J. Aubert: 3m 3f ( BYUC) ; Provence-Alpes-Côte d’Azur Region, Southern French Alps, Var, Artuby , La Martre, Pont Romain, 13.04.200 9, leg. Brulin: 1m ( GVC) ; Provence-Alpes-Côte d’Azur Region, Southern French Alps, Alpes-de-Haute-Provence, Esteron, Soleihas , 1100 m, 21.02.200 9, leg. Le Doare: 7m 9f ( GVC) ; Provence-Alpes-Côte d’Azur Region, Southern French Alps, Alpes-Maritimes, Lane , Andon , 1145 m, 13.04.200 9, leg. Le Doare: 2m 1f ( GVC) ; Franche-Comté Region, Jura, La Clauge, Belmont, Jura Forest Reserve , 10.04.200 9, leg. Le Doare: 2m 1f ( GVC) ; Limousin Region, French Massif Central, Creuse, Tarde , Champagnat , 440 m, 03.03.201 0, leg. Le Doare: 2m 1f ( GVC) ; Poitou-Charentes Region, French Massif Central, Vienne, Moulisme , Petite Blourde, N 147 bridge, 20.02.200 9, leg. Le Doare: 3m 4f ( GVC) ; Brittany Region, Bretagne, Finistère, Le Men, Tremorel , 105 m, 21.03.201 0, leg. Brulin: 5m 1f ( GVC) ; Brittany Region, Bretagne, Morbihan, Pont Drimo , Le Cours, 65 m, 12.02.201 1, leg. Brulin: 1m ( GVC) ; GERMANY: Schleswig-Holstein State, Plön, stream at Engelau , 26.02.196 6, leg. P. Zwick: 33m, 1m 2f larvae ( PZC) ; 03.196 6, leg. P. Zwick: 13m 2f, 3f larvae ( BYUC) ; Schleswig-Holstein State, Dallsek , 30 km E of Hennburg, 01.03.196 6, leg. P. Zwick: 18m ( BYUC) ; HUNGARY: Vas County, Őrség, Kercaszomor, Kerca Stream , 16.04.200 2, leg. A. Ambrus, T. Kovács: 2f ( MM) ; 11.03.200 3, leg. A. Ambrus, P. Juhász, T. Kovács: 1m 1f ( MM) ; Veszprém County, Bakony Mts., Döbrönte, Bittva Stream , 230 m, 30.03.200 6, leg. J. Kontschán, D. Murányi : 2m 1f ( HNHM) ; Veszprém County, Bakony Mts., Zirc- Szarvaskút , 21.04.198 8, leg. S. Tóth: 3m ( HNHM) ; Pest County, Pilis Mts., Pilisszentkereszt, Szurdok Stream , 25.03.200 6, leg. L. Dányi: 2m ( HNHM) ; Pest County, Börzsöny Mts., Szokolya, Les Valley , 08.02.200 3, leg. L. Papp, Zs. P. - Bajza: 3m 4f ( HNHM) ; 08.04.200 4, leg. D. Murányi : 1m, 4f larvae ( HNHM) ; Pest County, Börzsöny Mts., Szokolya-Királyrét, Bagoly-bükki Stream , 25.03.200 3, leg. T. Kovács: 1m ( MM) ; 18.03.200 4, leg. A. Ambrus: 3m 7f ( MM) ; Pest County, Börzsöny Mts., Szokolya-Királyrét, Szén Stream , 25.03.200 3, leg. T. Kovács: 1m ( MM) ; Pest County, Börzsöny Mts., Szokolya-Királyrét, Nagy-vasfazék Stream , 25.03.200 3, leg. T. Kovács: 1m ( MM) ; Pest County, Börzsöny Mts., Kismaros, Káposztás , Morgó Stream , 20.02.200 3, leg. P. Juhász, T. Kovács: 3m ( MM) ; Nógrád County, Cserhát Mts., Cserhátszentiván, Zsunyi Stream , 03.03.200 4, leg. K. Harmos: 3m ( HNHM) ; Nógrád County, Cserhát Mts., Ecseg, Zsunyi and Cserkúti Streams , 26.02.200 3, leg. K. Harmos: 9m 11f ( HNHM) ; Nógrád County, Cserhát Mts., Felsőtold, Szurdok Stream , 12.03.200 3, leg. K. Harmos: 8m 6f ( HNHM) ; Nógrád County, Cserhát Mts., Garáb, Garábi Stream , 04.03.200 3, leg. K. Harmos: 16m 5f, 1m 4f larvae ( HNHM) ; Nógrád County, Mátra Mts., Bátonyterenye, Lengyendi Stream at the forester house, 14.02.200 4, leg. K. Harmos: 1m ( HNHM) ; Heves County, Mátra Mts., Mátraszentimre, Csörgő Stream , 13.03.200 4, leg. K. Harmos: 2m, 1 f larva ( HNHM) ; Heves County, Mátra Mts., Mátrafüred, Bene Stream , 20.02.200 4, leg. D. Murányi : 1m ( HNHM) ; Heves County, Mátra Mts., Gyöngyössolymos, Cserkő-bánya , Monostor Stream , N 47°49.918’ E 19°55.801’, 260 m, 16.03.200 1, leg. T. Kovács: 3m 1f ( MM) ; 25.03.200 1, leg. T. Kovács: 3f ( MM) ; 19.04.200 1, leg. T. Kovács: 8f ( MM) ; 14.03.200 2, leg. T. Kovács: 3f, 3f exuviae ( MM) ; 19.04.200 2, leg. T. Kovács: 1f ( MM) ; 22.02.200 3, leg. T. Kovács: 1m 3f ( MM) ; Heves County, Mátra Mts., Gyöngyössolymos, Cserkő-bánya , Nagy Stream , 25.03.200 1, leg. T. Kovács: 1m 1f ( MM) ; 14.03.200 2, leg. T. Kovács: 7m ( MM) ; 08.03.200 3, leg. D. & T. Kovács: 2m ( MM) ; 31.03.200 4, leg. T. Kovács: 1m ( MM) ; 13.03.200 5, leg. T. Kovács: 2m ( MM) ; Heves County, Mátra Mts., Domoszló, Tarjánka Stream , 16.03.200 2, leg. D. Murányi : 3m ( HNHM) ; Heves County, Bükk Mts., Nagyvisnyó, Nagy-völgyi Stream , 22.01.200 3, leg. D. Murányi : 2m ( HNHM; one male used for drawings Figs. 141 , 154 ) ; Borsod-Abaúj-Zemplén County, Zemplén Mts., Vágáshuta, Hercegkúti Stream , 170 m, 14.03.200 4, leg. D. Murányi , Zs. Sóvári: 3m 1f, 1m larva ( HNHM) ; Borsod-Abaúj-Zemplén County, Zemplén Mts., Erdőhorváti, Tolcsva Stream at the third bridge, 170 m, 15.03.200 3, leg. D. Murányi , Zs. Sóvári: 4m, 2m larvae, 1m exuviae ( HNHM) ; ITALY: Lazio Region, Falscione Brooklet , 450m, 06.03.198 9, leg.?: 3m 2f ( BYUC) ; NORWAY: Akershūs County, Hūrdal, Fallbekken , 13.04.196 9, leg. Smedstad: 1m 1f ( BYUC) ; ROMANIA: Cluj County, Cluj- Napoca, Gorbo valley , 14.03.196 4, leg. B. Kis: 36m 26f ( HNHM) ; SLOVAKIA: Banskobystrický Region, Krupinská Planina, Čebovce, Čebovský Stream , 180 m, 24.03.200 6, leg. L. Dányi, J. Kontschán, D. Murányi : 1m, 3f larva ( HNHM) ; SWEDEN: Gotland County, SK Rövarkulan, 25.03.194 9, leg. P. Brinck: 2m 2f ( BYUC) ; UNITED KINGDOM: Scotland, Carluke , early 20 th century?, leg. K.J. Morton?: 1m ( WNHM) .

Description. Head, thorax, appendages and basal segments of the abdomen generotypic. Males micropterous, females macropterous. Dimensions of the presently examined specimens: body length: males 7.0–9.0, females 8.0–10.5 mm; forewing length: males 0.8–1.3 mm, females 8.0–9.5 mm.

Male terminalia ( Figs. 82–85 , 120 ): Process of Tg 9 high, perpendicularly elevated, its apex is less than one and a half times wider than the medial section of Ep-scl; its shape is rectangular, the apex smooth or bears two hardly visible hump-like tips; sides constantly narrow in caudal view, the membranous portion narrowest in the apical portion ( Figs. 152–156 ). Tg 10, B-scl and Lb-scl generotypic. Ep-scl wide and blunt in dorsal view, not swollen medially, medial width the ⅔ of basal width; tip upcurved in lateral view, divided section short. Ventral membranous part between the divisions of Ep-scl nearly reaches the base in lateral view; apical spines short, distributed only on the membranous part ( Figs. 106 , 114 , 139–143 ). I-scl generotypic, Ec short and often partly or fully everted on the non in-copula specimens. St 9 slightly projecting medially, vesicle large to medium sized, Fig. 83 shows the largest size of the range. Sg rounded with not so pronounced triangular shape, tip usually rounded. Pp, Fp, Rp and cerci generotypic.

Female subgenital plate ( Fig. 62 ): Rectangular, posterior margin slightly rounded and usually equal to the segment`s posterior margin. Antero-lateral recess usually distinct; the plate is entirely brown; lateral sclerites relatively large.

Drumming: Males produce repeated sequences of 2–3 drumming calls ( Fig. 165 ), but sporadically single calls can also be observed. Calls are monophasic: inter-beat intervals increase gradually during the beat sequence of a call ( Figs. 169–173 , 190 , Appendix Table 2 ). Calls in a sequence are rather similar to each other. The peak amplitude of beats generally shows a crescending-decrescending pattern during each call. See Figs. 169–173 for the oscillographic pattern of the male drumming calls of this species and also Table 7 for descriptive statistics of the examined five sonometric parameters. The male female drumming duet a long sequence of male call—female answer interchange ( Fig. 185 ).

Genetics: Three different haplotypes were found, two from Hungary and one from Serbia ( Fig. 192 ), with 0.5% of divergence, and with seven diagnostic characters separating them. Zwicknia bifrons specimens from Maljen Mts. of Serbia share haplotypes with Z. acuta . Despite this, morphology and mating calls correspond to Z. bifrons in all cases.

Affinities. Zwicknia bifrons is closely related to Z. acuta , and this affinity is discussed above. The epiproct is similar to Z. rupprechti , but the Tg 9 process is high and perpendicularly elevated in contrast to being low and caudally projecting in Z. rupprechti . In addition to Z. acuta and Z. rupprechti , males of Z. bifrons are easily distinguishable from other Zwicknia on the basis of upcurved Ep-scl tip and high, rectangular process of Tg 9. Females are difficult to distinguish, and the larvae are morphologically indistinguishable. The drumming signals of Z. bifrons are clearly different from those of the other three Zwicknia species where drumming was examined. This species produce the longest calls and drums with the slowest beat repetition rate among the investigated species. These two rhythmic characters clearly separate the calls of this species from the calls of the other three species which drum faster and produce shorter male calls (see Figs 190 , 191 ). Regarding the number of calls in a sequence, the male drumming signals of Z. bifrons are intermediate and overlap with Z. acuta and Z. rupprechti (and Z. kovacsi ). Z. bifrons differs genetically from the other species with 10 diagnostic characters and with> 2% nucleotide divergence.

Distribution and ecology. This species was originally described from Scotland. Drumming data has confirmed the occurrence of this species throughout the British Isles, Scandinavia, and southwards into the Alps ( Fig. 196 ). We have confirmed Z. bifrons in northern and Central Hungary. Specimens of this species have also been identified from Austria, France, Germany, and Romania on the basis of morphology ( Fig. 196–197 ). Adults were collected in January to May from the vicinity of slow to moderately fast flowing, medium-sized streams, usually in alder ( Alnus glutinosa ), but also in beech ( Fagus sylvatica ) forests between 150 and 600 m.

Remarks. The species was described on the basis of one or more females. A single female type apparently existed ( Morton 1896). Despite that females of Zwicknia are difficult to specifically distinguish, its identity with the above material studied is supported by the homogeneity of British populations, both in drumming signals ( Rupprecht 1997) and morphology based on the previously published figures (see under synonymies).

Figures

FIGURE 190. Inter-beat interval variation during male calls in the four examined Zwicknia Murányi, gen. n. species.—Box plot colors: white—Z. bifrons (Newman, 1838), red—Z. acuta Murányi & Orci, sp. n., blue—Z. rupprechti Murányi, Orci & Gamboa, sp. n. In Z. kovacsi Murányi & Gamboa, sp. n. we have drumming samples only from two specimens therefore in the case of that species two simple mark series are used to show inter-beat intervals in their calls (circles for specimen 1, and triangles for specimen 2). i1, i2, i3 and so on are labels containing the serial number of inter-beat intervals counting them from the beginning of each call. Box plots show the duration values measured in 7 specimens in Z. bifrons and Z. acuta, and 5 specimens in Z. rupprechti. Vertical axis has a logarithmic scale. Only calls recorded at an ambient air temperature of 17–19 oC were used. (see the measurement data in Appendix Tables 1–4).

FIGURE 191. A scatter plot showing the clear separation of male calls of Zwicknia Murányi, gen. n. along two drumming signal characters—Z. bifrons (Newman, 1838): triangles; Z. acuta Murányi & Orci, sp. n.: circles; Z. kovacsi Murányi & Gamboa, sp. n.: +; Z. rupprechti Murányi, Orci & Gamboa, sp. n.: X—Each mark represents the mean values of the characters for a given specimen. Note that both axes have logarithmic scale. Air temperature varied between 15–21 oC in every species (for details see Tables 6–8).

FIGURE 192. Phylogenetic tree of cox1 including Zwicknia Murányi, gen. n. with our accession codes (300941-301003), and GenBank closest species (with alphanumeric accession codes) as outgroup. MP and ML had same topology. Numbers on braches are ML/MP bootstrap values>50%. Species names right to the tree refer to morphological species. P-Distance table is showed.

FIGURES 1–6. Male terminalia of West Palaearctic Capniidae—1–2: Zwicknia bifrons (Newman); 3–4: Capnia s.s. nigra (Pictet); 5–6: Capnia s.l. vidua Klapálek; 1, 3, 5: dorso-caudal view; 2, 4, 6: ventro-caudal view (B-scl: basal sclerite; Cs: caudal setae; Ec: eversible crest; Ep-scl: main epiproct sclerite (shaded in grey); I-scl: inner sclerite; Lb-scl: laterobasal sclerite; Ll: lower limb; Tg 10: tergite 10)—not to scale.

FIGURES 7–10. Male genitalia of Zwicknia Murányi, gen. n.—7, 10: Z. bifrons (Newman); 8–9: Z. rupprechti Murányi, Orci & Gamboa, sp. n.; 7: Fp disconnected to the epiproct, latero-caudal view; 8: epiproct tip with contorted Ec, dorsal view; 9: epiproct with partly everted Ec and Fp connected to the Ep-scl, dorsal view; 10: epiproct with everted Ec, dorso-lateral view (B-scl: basal sclerite; Ec: eversible crest; Ep-scl: main epiproct sclerite; Fp: fusion plate; Lb-scl: laterobasal sclerite; Pp: paraproct; Sg: subgenital plate; Tg 9 p: process of tergite 9; Tg 10: tergite 10)—scales 0.1 mm.

FIGURES 23–31. Male paraprocts and fusion plate of Capniidae, dorso-lateral view—23: Zwicknia bifrons (Newman); 24: Capnia s.s. nigra (Pictet); 25: Capnia s.l. vidua Klapálek; 26: Capnia s.l. prolongata Zhiltzova; 27: Capnia s.l. pedestris Kimmins; 28: Capnia s.l. arensi Zhiltzova; 29: Bolshecapnia gregsoni (Ricker); 30: Isocapnia grandis (Banks); 31: Eucapnopsis brevicauda (Claassen) (Fp: fusion plate; Pp: paraproct; Rp: retractoral plate)—not to scale.

FIGURES 32–48. Male terminalia of Capniidae, ventral view—32: Zwicknia bifrons (Newman); 33: Bolshecapnia gregsoni (Ricker); 34: Eucapnopsis stigmatica transversa Aubert; 35: Isocapnia grandis (Banks); 36: Capnia s.l. arensi Zhiltzova; 37: C. s.l. prolongata; 38: C. s.l. pedestris Kimmins; 39: C. s.s. nigra (Pictet); 40: Arsapnia decepta Banks; 41: Allocapnia granulata (Claassen); 42: Mesocapnia silvatica Raušer; 43: Capnura venosa Banks; 44: Utacapnia lemoniana (Nebeker & Gaufin); 45: Paracapnia opis (Newman); 46: Nemocapnia carolina Banks; 47: Capnopsis schilleri schilleri (Rostock); 48: Capnioneura nemuroides Ris (C: cercus; Ep-scl: main epiproct sclerite; Fp: fusion plate; Pp: paraproct; Sg: subgenital plate; St 9: sternite 9; Tg 9: tergite 9; Tg 10: tergite 10; Vv: ventral vesicle)—scale 1 mm.

FIGURES 56–57. Macropterous wings of Zwicknia Murányi, gen. n.—56: Female Z. bifrons (Newman, 1838); 57: male Z. tuberculata (Zhiltzova, 1964) (A: anal vein; a: anal crossvein; arc: arculus; C: costa; Cu: cubital vein; cu-a: cubitoanal crossvein; h: humeral crossvein; M: medial vein; m-cu: mediocubital crossvein; R: radial vein; r: radial crossvein; r-m: radiomedial crossvein; Sc: subcosta)—scale 1 mm.

FIGURES 58–61. Micropterous wings of Zwicknia Murányi, gen. n. males—58: Z. bifrons (Newman, 1838); 59: Z. acuta Murányi & Orci, sp. n.; 60: Z. rupprechti Murányi, Orci & Gamboa, sp. n.; 61: Z. sevanica (Zhiltzova, 1964)—scale 0.2 mm.

FIGURES 62–65. Female genitalia of Zwicknia Murányi, gen. n.—62–63: Z. bifrons (Newman, 1838); 64: Z. acuta Murányi & Orci, sp. n.; 65: Z. rupprechti Murányi, Orci & Gamboa, sp. n.; 62: terminalia, ventral view; 63: inner genital organs, dorsal view; 64–65: subgenital plates, ventral view (gc: genital cavity; go: genital opening; Sg: subgenital plate)—scales 0.2 mm.

FIGURES 66–71. Larva of Zwicknia Murányi, gen. n.—66: Z. bifrons (Newman, 1838); 67-69, 71: Z. acuta Murányi & Orci, sp. n.; 70: Z. rupprechti Murányi, Orci & Gamboa, sp. n.; 66: head, dorso-lateral view; 67: mouthparts, ventral view; 68: pro- and mesothorax, ventral view; 69: meso- and metathorax, ventral view; 70: Tergite 5, dorsal view; 71: female terminalia, ventral view (Es: ecdysial suture; Etn: eutrochantin; Gl: glossa; Lb: labrum; Lp: labial palp; Me: mentum; MeFs: mesothoracic furcasternum; MeFsa: mesothoracic furcasternal arm; MeFsp: mesothoracic furcasternal pit; MeTn: mesothoracic trochantin; MFs: metathoracic furcasternum; Mp: maxillary palp; MTn: metathoracic trochantin; Oc: frontal ocellus; Pc: pedicell; PFs: prothoracic furcasternum; Pg: paraglossa; Pp: paraproct; St 8: Sternite 8)—scales 0.2 mm.

FIGURES 72–77. Larva of Zwicknia Murányi, gen. n.—72, 74, 76: Z. bifrons (Newman, 1838); 73, 75: Z. rupprechti Murányi, Orci & Gamboa, sp. n.; 77: Z. acuta Murányi & Orci, sp. n.; 72-73: hind femur, lateral view; 74-75: male terminalia, lateral view; 76-77: 11-13 cercal segments, lateral view (Pp: paraproct; Tg 10: tergite 10)— scales 0.2 mm.

FIGURES 82–85. Male terminalia of Zwicknia bifrons (Newman, 1838); Hungary, Pest County, Pilis Mts., Szentendre—82: dorsal view; 83: ventral view; 84: lateral view; 85: caudal view—scale 1 mm.

FIGURES 106–109. Lateral view of male epiproct of Zwicknia Murányi, gen. n.—106: Z. bifrons (Newman, 1838); 107: Z. rupprechti Murányi, Orci & Gamboa, sp. n.; 108–109: Z. acuta Murányi & Orci, sp. n.—scale 0.1 mm.

FIGURES 114–119. Dorsal view of male epiproct of Zwicknia Murányi, gen. n.—114: Z. bifrons (Newman, 1838); 115, 117: Z. rupprechti Murányi, Orci & Gamboa, sp. n.; 116: Z. kovacsi Murányi & Gamboa, sp. n.; 118: Z. sevanica (Zhiltzova, 1964); 119: Z. acuta Murányi & Orci, sp. n.—scale 0.1 mm.

FIGURES 120–123. Male terminalia of Zwicknia Murányi, gen. n., dorso-caudal view—120: Z. bifrons (Newman, 1838); 121: Z. acuta Murányi & Orci, sp. n.; 122: Z. rupprechti Murányi, Orci & Gamboa, sp. n.; 123: Z. kovacsi Murányi & Gamboa, sp. n.—scale 1 mm.

FIGURES 127–147. Male epiproct of Zwicknia Murányi, gen. n.—127–130 Z. acuta Murányi & Orci, sp. n.; 131–132: Z. kovacsi Murányi & Gamboa, sp. n.; 133–138: Z. rupprechti Murányi, Orci & Gamboa, sp. n.; 139–143: Z. bifrons (Newman, 1838); 144–145: Z. sevanica (Zhiltzova, 1964); 146: Z. tuberculata (Zhiltzova, 1964); 147: Z. turkestanica turkestanica (Kimmins, 1950); 127–131, 133, 135, 137, 139–147: lateral view; 132, 134, 136, 138: dorsal view (127: Slovakia, Krupinská Planina, Pôtor; 128: Hungary, Zemplén Mts., Kishuta; 129: Serbia, Fruska Gora, Vrdnik; 130: Serbia, Zlatibor Mts., Crni Rzav; 131–132: Romania, Rodna Mts., Borşa–Staţiunea Borşa; 133–134, 137–138: Croatia, Ivanščica Mts., Stari Golubovec; 135–136: Hungary, Mecsek Mts., Komló-Zobákpuszta; 139: Hungary, Börzsöny Mts., Szokolya-Királyrét; 140: Hungary, Mátra Mts., Gyöngyössolymos; 141: Hungary, Bükk Mts., Nagyvisnyó; 142: Hungary, Bakony Mts., Bakonybél; 143: Serbia, Maljen Mts., Brajkovići; 144–145: Turkey, Akyarma Pass; 146: Kazakhstan, Karatau Mts., Bajaldyrtal; 147: Kazakhstan, Tarbagatai Mts., Ayagos)—scale 0.2 mm.

FIGURES 148–164. Humps of male Tergite 9 of Zwicknia Murányi, gen. n., caudal view—148–151: Z. acuta Murányi & Orci, sp. n.; 152–156: Z. bifrons (Newman, 1838); 157–159: Z. rupprechti Murányi, Orci & Gamboa, sp. n.; 160: Z. kovacsi Murányi & Gamboa, sp. n.; 161–162: Z. sevanica (Zhiltzova, 1964); 163: Z. tuberculata (Zhiltzova, 1964); 164: Z. turkestanica turkestanica (Kimmins, 1950) (148: Slovakia, Krupinská Planina, Pôtor; 149: Hungary, Zemplén Mts., Kishuta; 150: Serbia, Fruska Gora, Vrdnik; 151: Serbia, Zlatibor Mts., Crni Rzav; 152: Hungary, Börzsöny Mts., Szokolya-Királyrét; 153: Hungary, Mátra Mts., Gyöngyössolymos; 154: Hungary, Bükk Mts., Nagyvisnyó; 155: Hungary, Bakony Mts., Bakonybél; 156: Serbia, Maljen Mts., Brajkovići; 157, 159: Croatia, Ivanščica Mts., Stari Golubovec; 158: Hungary, Mecsek Mts., Komló-Zobákpuszta; 160: Romania, Rodna Mts., Borşa-Staţiunea Borşa; 161–162: Turkey, Akyarma Pass; 163: Kazakhstan, Karatau Mts., Bajaldyrtal; 164: Kazakhstan, Tarbagatai Mts., Ayagos)—scale 0.2 mm.

FIGURES 165–168. Slow oscillograms of male calls of four Zwicknia Murányi, gen. n. species, showing call sequences versus sporadic call production—165: two call sequence of Z. bifrons (Newman, 1838), "//" indicates where 2 minutes 23 seconds of silence was cut out from the recording; 166: three call sequences of Z. acuta Murányi & Orci, sp. n.; 167: sporadic calls of Z. rupprechti Murányi, Orci & Gamboa, sp. n.; 168: sporadic call production in Z. kovacsi Murányi & Gamboa, sp. n.—time scale is the same for the four oscillograms; for detailed data on the animals and recording circumstances, see Table 9.

FIGURES 169–184. Male drumming calls of Zwicknia Murányi, gen. n.—169–173: Z. bifrons (Newman, 1838); 174–178: Z. acuta Murányi & Orci, sp. n.; 179–182: Z. rupprechti Murányi, Orci & Gamboa, sp. n.; 183–184: Z. kovacsi Murányi & Gamboa, sp. n. (see detailed data on the animals and recording circumstances in Table 9)—Oscillograms in Figs. 169–179, 181, 183 have the same time axis resolution (time scale bar is presented in Fig. 169 for all of them), an other time scale is used in the faster oscillograms of Figs. 180, 182, 184 (time scale bar for them can be found in Fig. 184), those are zoomed sections of Figs. 179, 181, 183, respectively.

FIGURES 185–189. Male-female drumming duets of three Zwicknia Murányi, gen. n. species.—185: Z. bifrons (Newman, 1838) from Hungary, Pilis Mts., Bükkös Stream, recorded at 16.9 °C; 186–187: Z. acuta Murányi & Orci, sp. n. from Slovakia, Krupinská Planina, Stará Rieka, recorded at 18.2 °C; 188: call-answer duet in Z. rupprechti Murányi, Orci & Gamboa, sp. n. from Hungary, Mecsek Mts., Völgységi Stream, recorded at 18.2 °C; 189: call-answer-reply duet in Z. rupprechti Murányi, Orci & Gamboa, sp. n. from Hungary, Mecsek Mts., Völgységi Stream, recorded at 17.9 °C—arrows with a gender sign point to the beginning of a section of the oscillogram containing the beats of that gender.

FIGURES 196–197. Distribution of Zwicknia Murányi, gen. n.—196: global distribution; 197: studied populations of the Carpathian Basin and the northern Balkans—in Fig. 196, white dot on Z. bifrons symbols indicate populations studied by Rupprecht (1997), dotted lines indicate limits of s.l. Z. bifrons literature reports; in Fig. 197, white dot on symbols indicate populations where drumming signals were studied by us, white star indicate type localities.