Zwicknia acuta Murányi & 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 : 37-40

Zwicknia acuta Murányi & Orci , sp. n.

( Figs. 59 , 64 , 67–69, 71 , 77 , 78–81 , 108–109 , 119 , 121 , 127–130 , 148–151 , 166 , 174–178 , 186–187 , 190–193 , 196–197 ).

Capnia nigra ( Pictet, 1833) —figures in Klapálek 1896 and Despax 1951 (enumerated under the general synonymies) possibly referring to Z. acuta .

Capnia bifrons ( Newman, 1838) —figures in Winkler 1957 (enumerated under the general synonymies) possibly referring to Z. acuta .

Diagnosis. Male epiproct: Ep-scl narrow and pointed in dorsal view, tip straight, long and acute in lateral view; ventral membranous section ends far before the base in lateral view, apical spines stout, distributed also on the apex of Ep-scl. Process of male Tg 9: high, perpendicularly elevated, 2× wider than Ep-scl, rectangular and with indenting sides caudally. Males produce sequences of (2)-3-4-(6) drumming calls ( Figs. 166 , 174–178 ). Calls are monophasic beat series in which beats are produced with gradually increasing inter-beat intervals ( Fig 190 , red box plots). Inter beat intervals vary between 70–130 ms. Call duration varies between 350–750 ms and calls contain 4–10 beats. Inter call intervals increase during call sequence and vary between 350–3250 ms. All these parameters refer to signals produced at ambient air temperature between 15–21 o C. Male female drumming duet is a long sequence of male call—female answer alternations, where female answers may overlap the terminal part of male calls ( Figs 186–187 ).

Type material. Holotype male: SLOVAKIA: Banskobystrický Region, Krupinská Planina, Pôtor, Stará Rieka Stream, N 48°13.731’ E 19°24.946’, 200 m a.s.l., 16.03.201 0, leg. D. Murányi , K. Orci ( HNHM: PLP3882; drumming recorded as 2010/No.1). Paratypes: same locality and date: 1f ( HNHM: PLP3883; drumming recorded as 2010/No.1, pair of the holotype), 1m ( HNHM: PLP3884; drumming recorded as 2010/No.2), 1m ( HNHM: PLP3885; drumming recorded as 2010/No.3), 1m ( HNHM: PLP3886; drumming recorded as 2010/No.4, Fig. 175 ), 1m ( HNHM: PLP3887; drumming recorded as 2010/No.5), 1m ( HNHM: PLP3888; drumming recorded as 2010/ No.6), 15m, 1f larva ( HNHM: PLP3372), 4m ( BYUC); 24.03.200 6, leg. L. Dányi, J. Kontschán, D. Murányi : 6m 2f, 1f larva ( HNHM: PLP1832; two male, two male terminalia, one female and the larva prepared for SEM, specimens used for drawings and photos Figs. 59 , 64 , 67–69, 71 , 77 , 78–81 , 108–109 , 119 , 121 , 127 , 148 ), 2m ( GVC), 2m ( PZC); 23.03.200 9, leg. N. Hordós, D. Murányi , J. Papp, Zs. Ujvári: 2m 2f, 1f larva ( HNHM: PLP3044); 23.03.201 1, leg. K. Orci: 1m ( HNHM: PLP3811; used for molecular studies as 300985, drumming recorded as 2011/No.1), 1m ( HNHM: PLP3812; used for molecular studies as 300982, drumming recorded as 2011/No.2), 1m ( HNHM: PLP3813; drumming recorded as 2011/No.3), 1m ( HNHM: PLP3814; used for molecular studies as 300992, drumming recorded as 2011/No.4), 1m ( HNHM: PLP3815; used for molecular studies as 301003, drumming recorded as 2011/No.5), 1m ( HNHM: PLP3816; drumming recorded as 2011/No.6), 1m ( HNHM: PLP3808; used for molecular studies as 300983, drumming recorded as 2011/No.7), 1m ( HNHM: PLP3881; used for molecular studies as 300981, drumming recorded as 2011/No.8), 1m ( HNHM: PLP3817); HUNGARY: Borsod-Abaúj-Zemplén County, Zemplén Mts., Kishuta, Kemence Stream, N 48°27.298’ E 21°28.707’, 175 m, 08.03.201 1, leg. T. Kovács, D. Murányi , K. M. Orci, G. Puskás: 3m ( HNHM: PLH1242; one male used for drawings Figs. 128 , 149 ), 1m ( HNHM: PLH1258; used for molecular studies as 300989, drumming recorded as 2011/No.1), 2m ( HNHM: PLH1300; used for molecular studies as 30100 and 30101, drumming not recorded), 1m (only bioacoustic study, specimen escaped; drumming recorded as 2011/No.4, Fig. 174 ); SERBIA: Syrmia District, Fruska Gora, Vrdnik, stream NW of the village, N 45°08.583’ E 19°46.299’, 275 m, 15.03.201 1, leg. T. Kovács, G. Magos, D. Murányi : 1m ( HNHM: PLP3818; used for drawings Figs. 129 , 150 , for molecular studies as 300996, drumming recorded as 2011/No.1, Fig. 176 ), 1m ( HNHM: PLP3889; used for molecular studies as 300962); Braničevo District, Homoljske Planina, Krepoljin, stream N of the village, N 44°16.582’ E 21°36.553’, 290 m, 16.03.201 1, leg. T. Kovács, G. Magos, D. Murányi : 1m ( HNHM: PLP3820; used for molecular studies as 300965, drumming recorded as 2011/No.1, Fig. 177 ), 1m ( HNHM: PLP3821; used for molecular studies as 300951); Zlatibor District, Zlatibor Mts., Crni Rzav Stream along the road No.21, N 43°39.731’ E 19°42.575’, 1010 m, 17.03.201 1, leg. T. Kovács, G. Magos, D. Murányi : 1m ( HNHM: PLP3582; used for drawings Figs. 130 , 151 ), 1m ( HNHM: PLP3819; used for molecular studies as 300994, drumming recorded as 2011/No.1), 1f ( HNHM: PLP3890; used for molecular studies as 300952), 1m (only bioacoustic study, specimen escaped; drumming recorded as 2011/No.3, Fig. 178 ).

Other material—Records based on morphology: AUSTRIA: Lower Austria State, Purgstal, Bahnhof, im Flug , 18.03.197 1, leg. Ressl: 1m ( PZC) ; only as ‘ Austria’ , 1861, leg. Roghf.: 2m ( WNHM) ; GERMANY: Schleswig-Holstein State, Busdorf , Kr. Schleswig, 15.01.195 1, leg. H. Dittmar: 2m ( PZC) ; Saxony-Anhalt State, Harz, Nagelbach , 15.03.200 5, Lutz Tappenbeck: 3m 1f ( PZC) ; HUNGARY: Nógrád County, Cserhát Mts., Nógrádszakál, Ipoly River at the gorge of Párizs Stream, 03.04.200 5, leg. T. Kovács: 1m ( MM) ; Borsod-Abaúj- Zemplén County, Zemplén Mts., Kishuta, Komlóska Stream , 220 m, 14.03.200 4, leg. D. Murányi , Zs. Sóvári: 1m ( HNHM) ; Borsod-Abaúj-Zemplén County, Zemplén Mts., Kishuta, Kemence Stream , 175 m, 14.03.200 4, leg. D. Murányi , Zs. Sóvári: 1f larva ( HNHM) ; SLOVAKIA: Banskobistrický Region, Javorie Mts., Horný Tisovník , Kostolné , Tisovník Stream , 470 m, 24.03.200 6, leg. L. Dányi, J. Kontschán, D. Murányi : 8m 1f ( HNHM) ; further 20 males present in the WNHM, collected during second half of the 19 th century, but without any reliable labels (some are possibly from Mainz or Bergen, collected by F. Brauer in 1865) .

Description. Head, thorax, appendages and basal segments of the abdomen generotypic. Males micropterous, females macropterous. Body length: holotype 7.5 mm, male paratypes 6.5–8.0, female paratypes 8.0–10.5 mm; forewing length: holotype 1.6 mm, male paratypes 1.3–1.7 mm, female paratypes 8.5–11.0 mm.

Male terminalia ( Figs. 78–81 , 121 ): Process of Tg 9 high, perpendicularly elevated, apex 2× wider than the medial section of Ep-scl; rectangular in shape, apex bearing two small hump-like tips; sides indented in caudal view, forming the membranous portion narrowest medially ( Figs. 148–151 ). Tg 10, B-scl and Lb-scl generotypic. Ep-scl narrow and pointed in dorsal view, medially not swollen, its medial width ½ to ⅓ of basal width; tip straight and acute in lateral view, divided section long. Ventral membranous part between the division of Ep-scl ends far before the base in lateral view; apical spines stout, distributed not only on the membranous part but extend to the Ep-scl ( Figs. 108–109 , 119 , 127–130 ). I-scl generotypic, Ec long and rarely everted on the non in-copula specimens. St 9 slightly projecting medially, vesicle medium sized to large, Fig. 79 illustrates the smallest of the range. Sg rounded with pronounced triangular shape, tip usually incised. Pp, Fp, Rp and cerci generotypic.

Female subgenital plate ( Fig. 64 ): Less rectangular than usual, posterior margin rounded and sometimes slightly overhanging the segment. Antero-lateral recess usually distinct, the plate is entirely brown; lateral sclerites relatively large.

Drumming: Males produce (2)-3-4-(6) drumming calls in a sequence ( Fig. 166 ). Calls in a sequence are similar to each other. However, the first call is generally shorter and of lower amplitude than the following ones. Calls are monophasic signals consisting of one beat group with inter beat intervals increasing gradually toward the end of beat group ( Fig 190 : red box plots, Appendix Table 1 ). The amplitude envelope of calls are of the following main types: a full length crescending (e.g. see the second call in Fig. 175 ), an initial crescendo+nearly constant main part (see 4 th call in Fig. 176 ) or a crescending first half+decrescending second half pattern (e.g. 4 th call in Fig. 177 ). Calls follow each other with increasing inter-call intervals. See Figs. 174–178 for the oscillographic pattern of the male drumming calls of this species. Descriptive statistics of five drumming characters are presented in Table 6 . See Fig. 190 for a box plot series showing the variation of inter-beat intervals within a call in relation to the other three Zwicknia species , where drumming signals are characterized in this study. Appendix Table 1 contains the measurement data of inter-beat intervals of this species. The male female drumming duet is a long sequence of male call—female answer, where the female answers sometimes overlaps the terminal part of male calls ( Figs. 186–187 ).

Genetics: Three different haplotypes were found corresponding well to the geographical distribution ( Hungary, Serbia, and Slovakia). The Slovakia haplotypes clustered at the same node as the northern Hungarian population, resulting in 0.4% nucleotide divergence and two diagnostic characters. We found that the specimens from Maljen Mts. of Serbia included individuals of both Z. bifrons and Z. acuta ( Fig. 192 ). However, despite that the morphology and mating calls correspond to Z. bifrons in all cases.

Affinities. This species is morphologically very close to Z. bifrons . One of the Serbian populations (Maljen Mts.) may be hybrids, as they share mitochondrial haplotypes ( Fig. 192 ). Individuals have drumming signals and morphology similar to Z. bifrons and illustrations are provided ( Figs. 143 , 156 , 173 ). However, nearby Slovakian and Hungarian populations not sympatric with Z. bifrons populations and other Serbian populations exhibit differences in both morphology and mating calls. Morphologically, the males differ by having narrow and pointed Ep-scl with an acute and straight apex, instead of wide and blunt Ep-scl and upcurved tip; ventral membranous section terminate far before the base instead of nearly reaching it, and Tg 9 process with indented sides caudally instead of consistently narrow sides. Males of Z. acuta are easily distinguishable from other Zwicknia species on the basis of acute Ep-scl tip and very high, rectangular process of Tg 9. Females are difficult to separate with certainty and the larvae are morphologically indistinguishable. The drumming signals of Z. acuta clearly differ from those of the other three Zwicknia species analysed ( Figs. 165–184 ). Male drumming calls are shorter ( Fig. 191 , Tables 6–8 ) and contain generally a fewer number of beats than in Z. bifrons , but are much longer than in Z. kovacsi and Z. rupprechti . The number of calls in a call sequence is generally higher, and calls are repeated with shorter inter call intervals than in Z. bifrons ( Figs. 165–166 , Tables 6–7 ). Molecular diagnostics of this species also yielded differences among the other species with a node of four diagnostic characters as compared to Z. bifrons and 1–5% of total divergence.

Distribution and ecology. The species is found in southern Slovakia, northeastern Hungary, northern and central Serbia. Specimens from Austria and Germany are attributed to this species on the basis of morphology ( Figs. 196–197 ). Adults were found in March emerging from slow or moderately fast flowing, large to mediumsized streams, mostly in alder ( Alnus glutinosa Gaertin. ) forests, but also in open grasslands between 200– 1,000m ( Fig. 193 ).

Etymology. The name acuta (from the latin word acutus, meaning acute) refers to the acute tip of the epiproct. Used as an adjective, gender feminine.

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 193–195. Type localities of the new species described in Zwicknia Murányi, gen. n.—193: Zwicknia acuta Murányi & Orci, sp. n.: Stará rieka, Slovakia, Banskobystrický Region, Krupinská Planina, Pôtor; 194: Zwicknia kovacsi Murányi & Gamboa, sp. n.: Cimpoies Stream, Romania, Maramureş County, Rodna Mts., Borşa-Staţiunea Borşa; 195: Zwicknia rupprechti Murányi, Orci & Gamboa, sp. n.: Völgységi Stream, Hungary, Baranya County, Mecsek Mts., Komló-Zobákpuszta.

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 78–81. Male terminalia of Zwicknia acuta Murányi & Orci, sp. n.; paratype, Slovakia, Banskobystrický Region, Krupinská Planina, Pôtor—78: dorsal view; 79: ventral view; 80: lateral view; 81: caudal view (Fp: fusion plate; Pp: paraproct)—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.