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) Author Murányi, Dávid muranyi@zool.nhmus.hu Author Gamboa, Maribet maribetg@gmail.com Author Orci, Kirill Márk muranyi@zool.nhmus.hu text Zootaxa 2014 2014-06-06 3812 1 1 82 journal article 5365 10.11646/zootaxa.3812.1.1 fd5ba21e-09ce-4ac6-b84f-56632ed93917 1175-5326 4919079 7847D731-9F66-4856-A79F-9435FED25B1D Zwicknia rupprechti Murányi, Orci & Gamboa , sp. n. ( Figs. 8–9 , 60 , 65 , 70 , 73, 75 , 90–93 , 107 , 115, 117 , 122 , 133–138 , 157–159 , 167 , 179–182 , 188–192 , 195–197 ) Capnia bifrons ( Newman, 1838 ) —drumming signals in Rupprecht 1982 ( Fig. 2c ) probably refers to Z. rupprechti . Capnia bifrons ( Newman, 1838 ) Capbif race sensu Rupprecht 1997 — Rupprecht 1997: 94 . (drumming signals, probably refers to Z. rupprechti ); Enting & Rupprecht 2001: 71 (clarification as not 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: very low, caudally projecting, <1½× wider than Ep-scl, rounded and with continuously narrow sides caudally. Male drumming calls are short beat groups with 60–80 ms duration and including 4–6 percussive beats produced with a mean inter beat interval of 17–23 ms. Inter-beat intervals gradually decrease, while the amplitude of beats noticeably increase during a call. Sometimes two calls follow each other with an inter- call interval of 1.5–5.5 seconds, but generally single calls are produced sporadically. The male-female drumming duet consists of a single male call—female answer or male call—female answer—male response signal exchanges as in Figs. 188–189 . Type material. Holotype male: HUNGARY : Baranya County , Mecsek Mts. , Komló-Zobákpuszta , Völgységi Stream , N 46°11.642’ E 18°19.071’ , 305 m a.s.l. , 09.03.201 0, leg. D. Murányi , K. M. Orci ( HNHM : PLH1277 ; drumming recorded as duett 2010/No.6) . Paratypes : same locality and date: 6m , 2m 2f larvae ( HNHM : PLH1184 ; three male terminalias, one female and one male larva prepared for SEM, specimens used for drawings and photos Figs. 8 , 70 , 73, 75 , 115 , 135–136 , 158 ), 1m 1f ( HNHM : PLH1274 ; drumming recorded as 2010/No.8), 1f ( HNHM : PLH1275 ; drumming recorded as 2010/No.2, male escaped), 1m 1f ( HNHM : PLH1276 ; used for molecular studies as 300970, 300971, drummings recorded as 2010/No.1), 1f ( HNHM : PLH1278 ; drumming recorded as duett 2010/ No.6, pair of the holotype ), 1m 1f ( HNHM : PLH1279 ; drumming recorded as 2010/No.5), 1m 1f ( HNHM : PLH1280 ; drumming recorded as 2010/No.7, Figs. 179–180 ), 3m ( BYUC ) , 1m 1f ( HNHM : PLH1297 ; used for molecular studies as 300948, 300949, drummings recorded as 2010/No.3), 1m 1f ( HNHM : PLH1298 ; used for molecular studies as 300963, 300964, drummings recorded as 2010/No.4), 1f ( HNHM : PLH1299 ; used for molecular studies as 300967, drumming recorded as 2010/No.9); CROATIA : Krapina-Zagorje County , Ivanščica Mts. , Stari Golubovec , Reka Stream below the village, N 46°10.54’ E 16°02.99’ , 345 m , 06.04.201 0, leg. L. Czigány , D. Murányi : 7m ( HNHM : PLP3401 ; one male prepared for SEM, specimens used for drawings and photos Figs. 60 , 90–93 , 122 , 133–134, 137–138 , 157, 159 ), 1m 1f ( HNHM : PLP3878 ; used for molecular studies as 300973, 300974, drummings recorded as 2010/No.2, Figs. 181–182 ), 1m ( HNHM : PLP3879 ; used for molecular studies as 300968, drumming recorded as 2010/No.1), 1m ( HNHM : PLP3880 ; drumming recorded as 2010/No.3), 2m ( GVC ) , 2m ( PZC ) . Other material—Records based on morphology: CROATIA : Krapina-Zagorje County , Ivanščica Mts. , Lobor , Reka Stream above the village, N 46°09’ E 16°05’ , 330 m , 01.04.200 6, leg. J. Kontschán , D. Murányi : 5m 1f ( HNHM ; one male terminalia prepared for SEM, used for photos Figs. 9 , 107 , 117 ) ; HUNGARY : Baranya County , Mecsek Mts. , Komló , Hidas Valley , 360 m , 28.02.200 4, leg. D. Murányi : 3m , 1f larva ( HNHM ) ; Baranya County , Mecsek Mts ., Pécs , Éger Valley , 250 m , 29.02.200 4, leg. D. Murányi : 1m 2f, 5m 6f larvae, 2m exuviae ( HNHM ; one female prepared for SEM Fig. 65 ) . Description. Head, thorax, appendages and basal segments of the abdomen generotypic. Occipital rugosities present on teneral specimens. Males micropterous, females macropterous. Body length: holotype 6.5 mm, male paratypes 6.0–6.5, female paratypes 8.0–8.5 mm; forewing length: holotype 1.6 mm, male paratypes 1.3–2.0 mm, female paratypes 8.0–8.5 mm. Male terminalia ( Figs. 90–93 , 122 ): Process of Tg 9 very low, caudally projecting, its apex is,1½ × wider than the medial section of Ep-scl; its shape is flattened elliptical, the apex smooth, rounded; sides continuously narrow in caudal view, the membranous portion narrowest apically ( Figs. 157–159 ). Tg 10, B-scl and Lb-scl generotypic. Ep-scl wide and blunt in dorsal view, medially not swollen, its medial width ⅔ basal width; tip upcurved in lateral view, divided section short. Ventral membranous part between the division of Ep-scl nearly reach the base in lateral view; apical spines short, distributed only on the membranous part ( Figs. 107 , 115, 117 , 133–138 ). I-scl generotypic, Ec short and often partly or fully everted on the non in-copula specimens, even on tenerals ( Figs. 133–138 ). St 9 slightly projecting medially, vesicle small to medium-sized, Fig. 91 illustrates the smallest size range. Sg rounded with pronounced triangular shape, tip usually rounded. Pp, Fp, Rp and cerci generotypic. Female subgenital plate ( Fig. 65 ): Rectangular, posterior margin slightly rounded, usually slightly incised and equal to the segment`s posterior margin. Antero-lateral recess usually distinct, the plate is entirely brown; lateral sclerites relatively large. Drumming: Males produce generally single, monophasic calls in a sporadic pattern. Within each call inter-beat intervals decrease gradually ( Fig 190 , Appendix Table 3 ). Peak amplitude of beats generally increases along the beat sequence of a call. See Figs. 179–182 for the oscillographic pattern of the male drumming calls of this species, and also Table 8 for descriptive statistics of the examined three sonometric parameters. The male-female drumming duet consists of a single male call—female answer signal exchanges, but also male call—female answer—male response intersexual signalling can be observed ( Fig. 189 .) Female answer signals are monophasic beat sequences of a similar beat repetition rate as observable in the male call, but are longer and contain more beats than the male signals. For a more detailed quantitative analysis of the male-female duet pattern, recordings from more females will be necessary. FIGURES 90–93. Male terminalia of Zwicknia rupprechti Murányi, Orci & Gamboa , sp. n. ; paratype, Croatia, Krapina- Zagorje County, Ivanščica Mts., Stari Golubovec — 90: dorsal view; 91: ventral view; 92: lateral view; 93: caudal view—scale 1 mm. TABLE 7. Descriptive statistics for the examined rhythmic characters of the male drumming calls of Zwicknia bifrons ( Newman, 1838 ) , and also for the ambient air temperature during the sound recordings.—"n" is the number of male specimens examined. In each character data points where mean values obtained for each examined male on the basis of 1-10 signal measurements per specimen. Number of examined specimens per population: Hungary: Pilis Mts— 7, Bakony Mts—2, Börzsöny Mts—4, Mátra Mts—5;Serbia: Maljen Mts—7.

n	mean	SD	min	max
Call duration (ms)	25	1811	264.7	1443	2389
Number of beats per call	25	11.26	1.684	7.33	15.5
Mean beat inteval (ms)	25	178.1	19.2	147.6	229
Inter-call interval within a call sequence (ms)	21	3061	1411.1	1447	6295
Number of calls in a call sequence	24	1.992	0.575	1	3
Air temperature (o C)	25	18.5	1.407	15.8	20.8

Genetics: Two different haplotypes from southwestern Hungary and northern Croatia clustered on the same node with 1% divergence with 5 informative characters separating them. All individuals collected in Hungary and Croatia clustered together. Affinities. The relationship of Z. rupprechti to Z. bifrons is discussed above. In addition, Z. rupprechti males are easily distinguishable from other Zwicknia on the basis of upcurved Ep-scl tip and low, caudally projecting process of Tg 9. Females cannot be distinguished from other species of the genus as are larvae morphologically indistinguishable. The male drumming calls of Z. rupprechti are of short duration and have a fast beat repetition, differing from those of Z. bifrons and Z. acuta , but similar to the calls of Z. kovacsi . Distinctive characters of the male calls of Z. rupprechti that differ from those of Z. kovacsi are shorter call durations, inter-beat intervals exhibiting a gradual decrease, and the beat peak amplitude with a characteristic increase during the beat call sequence. The calls of Z. kovacsi exhibit less of a change in those signal parameters during a call. With their short mean inter-beat intervals the calls of Z. rupprechti are closer to Rupprecht's "Capbif" call variant than the slightly slower calls of Z. kovacsi , which stand a little farther form "Capbif". However, regarding the number of beats per call, it is Z. kovacsi that is closer to "Capbif" (compare our results to the "Capbif" drumming data in Rupprecht 1997 ). Male call differences are minor and to examine their significance in female mate choice will require additional studies including play-back experiments. However, the male terminalia of Z. rupprechti is similar to populations producing "Capbif" type signals and Z. kovacsi exhibits clear distinctive features from them. Zwicknia rupprechti differs from other species of the genus by 8% genetic divergence with 17 informative characters. Distribution and ecology. Zwicknia rupprechti occurs in southwestern Hungary and northern Croatia ( Figs. 196–197 ). Populations from the southern Alps have similar drumming signals ( Rupprecht 1997 ), but are not considered to be this species because of the close similarities of the drumming calls with those of Z. kovacsi . Adults of Z. rupprechti are active from March to early April and are associated with slow or moderately fast, mediumsized streams flowing through alder ( Alnus glutinosa ) forests between 250 and 350 m ( Fig. 195 ). Etymology. The species is dedicated to Dr. Rainer Rupprecht, Mainz, Germany , in recognition of his primary contribution to our knowledge on Plecoptera drumming calls; furthermore, he recognized the distinctiveness of the present drumming type . Used as the genitive of a noun of male gender.