Zwicknia Murányi, 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 : 24-37

Zwicknia Murányi , gen. n.

( Figs. 1–2 , 7–10 , 23 , 32 , 49–197 )

Capnia nigra ( Pictet, 1833) sensu Morton 1896 — Morton 1896: 60. (complementary description of the adult); Klapálek 1896: 19. (detailed description of the male genitalia); Klapálek 1909: 55, 88 (complementary description of the adult, description of the larva); Hynes 1940: 18. (complementary description of the adult); Hynes 1941: 498. (complementary description of the larva); Hanson 1946: 194. (detailed morphology of the adult); Despax 1951: 154. (complementary description of the adult).

Capnia quadrangularis Aubert, 1946 — Aubert 1946: 24. (complementary description of the larva).

Capnia bifrons ( Newman, 1838) — Brinck 1949: 96. (complementary description of the larva); Kimmins 1950a: 188. (complementary description of the adult); Kimmins 1950b: 9. (complementary description of the adult); Aubert 1951: 281. (complementary description of the larva); Brinck 1952: 56, 111. (complementary description of the adult and the larva); Hynes 1955a: 92. (complementary description of the larva); Hynes 1958: 37, 67. (revised complementary descriptions from Hynes 1940 and 1955a; same in the further editions); Illies 1955: 78. (complementary description of the adult); Winkler 1957: 43. (complementary description of the adult); Aubert 1959: 73, 125. (complementary description of the adult and the larva); Khoo 1964: 29. (description of the aestivating larva); Lillehammer 1965: 49. (complementary description of the adult and the larva); Rupprecht 1965: 1258. (groundplan of mating call); Rupprecht 1968: 43. (details of mating call); Zwick 1973: 163, 183. (terminal morphology of the male); Kis 1974: 119. (complementary description of the adult); Lillehammer 1974: 92. (variability of the adult); Rupprecht 1976: 38. (terminal morphology and different drumming signals of the male); Rupprecht 1982: 96. (different drumming signals of the male); Lillehammer 1988: 136. (complementary description of the adult and the larva); Westermann 1993: 137. (wing polymorphy); Rupprecht 1997: 94. (different drumming signals); Tierno de Figueroa et al. 2003: 242. (complementary description of the adult); Zwick 2004: 320. (further larval characters); Fochetti & Tierno de Figueroa 2008: 227. (complementary description of the adult).

Capnia bifrons species group sensu Zhiltzova 2001 — Zhiltzova 2001: 424. (definition); Zhiltzova 2003: 345. (revision).

Diagnosis. Male epiproct: B-scl large, divided from Ep-scl; Lb-scl small, divided from Ep-scl; Ep-scl ventrally fused at base and tip, laterally divided in the apex, caudal setae absent; I-scl long, open tube, Ec present. Male Pp: apical part short and wide; Fp long and narrow, divided from Rp. Male Sg: divided from St 9 and Tg 9, vesicle present. Female Sg: rectangular, entire, small lateral sclerites present. Male tergites: Tg 9 with process. Ventral thoracic sclerites: MPrs and MeFs triangular, MeFsp separated from MePfs. Macropterous wings: forewing A1 beyond a and R1 before r curved.

Type species. Zwicknia bifrons ( Newman, 1838) = Chloroperla bifrons Newman, 1838 .

Further species included. Zwicknia acuta Murányi & Orci , sp. n., Zwicknia kovacsi Murányi & Gamboa , sp. n., Zwicknia rupprechti Murányi, Orci & Gamboa , sp. n., Zwicknia sevanica ( Zhiltzova, 1964) comb. n., Zwicknia tuberculata ( Zhiltzova, 1964) comb. n., Zwicknia turkestanica turkestanica ( Kimmins, 1950a) comb. n., Zwicknia turkestanica brevicula ( Berthélemy & Dia, 1982) comb. n.

Description. Medium sized Capniidae , males usually micropterous, also brachypterous or macropterous, females always macropterous. Body length related to season of emergence, smallest late males are 4.5 mm while largest early males are 10.0 mm; females up to 12.0 mm. Pilosity generally short and dense, longer setae occur on the appendages, especially on cerci and femora; male epiproct bare beside B-scl, pilosity of Tg 9 process reduced. Fully sclerotized adults dark brown to black, with even darker rugosities on head and pronotum; wings hyaline, venation dark brown ( Figs. 49–51 ).

Teneral coloration ( Fig. 52 ): Mouthparts, palpi and antennae pale, eyes black. Head capsule pale, tentorial callosities and enlarged M-line dark; occipital rugosities usually lacking on teneral specimens. Pronotum pale with brown areas extending around the dark rugosities, central line, front and posterior delimiting folds of pronotal sclerite. Meso- and metanotum of the male with dark Scl and inner corners of Pra, Sct with paired lateral dark patches and a darker stripe beyond Scl. Meso- and metanotum of the female mostly pale with dark inner corners of Pra and limits of the Scl. Wings whitish with pale veins in both sexes. Laterally the thorax is pale with parts of Epm and Tn or Etn dark; legs pale but dorsal part of coxae dark. Ventral surface of the thorax is pale with dark Fs and Fsp. Abdominal tergites of the male with dark brown antecosta, transverse row of four spots present on Tg 1–9. Tg 1 mostly brown, extent of brown color declining on the segments towards the terminalia but gradually darken anteriorly and laterally on the tergites. Process of Tg 9 pale, Tg 10 entirely pale but with two lateral spots. B-scl pale but with dark anterior margin, Lb-scl entirely pale, Ep-scl entirely dark. Abdominal tergites of the female mostly pale, Tg 1–9 laterally with paired longitudinal patch and a pair of spots; Tg 10 entirely pale. Abdominal sternites of the male mostly pale, St 3–8 with anterior sclerites forming a medially interrupted, dark transverse line; Sg with a paired anterolateral spot, vesicle pale. Pp pale with brown tip, cerci pale. Ventral surface of the female abdomen entirely pale.

Head: Submentum and mentum small, general Capniidae shaped; glossae apically rounded, paraglossae weakly subdivided, labial palpi short with second palpomere the longest. Maxillae and other mouthparts are of general Capniidae shape, maxillary palpi moderately long with the third palpomere the longest. Head shape usual of Capniidae , eyes smaller than the area delimited by the three ocelli. Ocelli and tentorial callosities prominent in fully sclerotized adults, ecdysial suture, M-line and occipital rugosities usually distinct. Antennae long, with 30 or more antennomeres. Antennomeres club or pearl-like shaped, gradually elongated towards the apex; some of the basal 4–6 antennomeres usually subdivided.

Thorax ( Figs. 53–55 ): Prothorax having large Pn with rounded corners. PPrs small, caudally projecting, not fused with the heart-shaped PBs; PPr narrow, fused with PBs; PFs transverse, fused with both PBS and the stripelike, curved PPo; PPfs large, elliptical, not fused with PFs. PEps and PEpm small, fused with each other and Peps to PPr, PEpm touch but not fused with the narrow, curved PEtn. Mesothorax with large MeSct surrounding MeScl, fused with the rectangular, longitudinally divided MePsc and then the wide, laterally subdivided MePra; MeTgl and subalar sclerites are small, none of them fused with other sclerites but the base of forewing; MePscl entire and fused with MeSct in females and macropterous males, while longitudinally divided and not fused with MeSct in micropterous males. MeSs very narrow, touch but not fused with PPfs while fused with the large, oblong MeBs; MePrs elliptical, not fused with MeBs; MeFs triangular and fused with MeBs, and with the distinct, paired MeFsa and the MeFsp; MePfs divided in two lateral, rounded parts by the MeFsp, but the parts are not fused with other sclerites. MeKes fused with MeBs and entirely fused with the ventrally elongated and narrowing MeTn; MeAes fused with MeKes, and projecting in elongated, hardly separated MeAb, MePb and MeAl, among which MePb ends in a globular apex; MeEpm ventrally fused with MeTn, MeKes, MeAes and MeAl, in females and macropterous males it is dorsally fused with MePscl while separated from in micropterous males. Metathorax with MSct smaller than MeSct, but as well surrounding MScl, fused with the rectangular, longitudinally divided MPsc, while MPsc hardly fused with the smaller MPra; MTgl and subalar sclerites are small, none of them fused with other sclerites but the base of hindwing; MPscl entire and fused with MeSct in females and macropterous males, while longitudinally subdivided and hardly fused with MeSct in micropterous males; MPscl fused with Tg 1 in both cases. MPrs triangular, not fused with the large, oblong MBs; MSs small and fused with MBs; MFs stripe-like and fused with MBs, laterally projecting backwards but not fused with St 1. MKes fused with MBs and entirely fused with the ventrally elongated and narrowing MTn; MAes fused with MKes, and projecting in elongated, hardly separated MAb, MPb and MAl, among which MPb ends in a globular apex; MEpm ventrally fused with MTn, MKes, MAes and MAl, in females and macropterous males it is dorsally fused with MPscl while separated from in micropterous males.

Legs: Medium sized in relation of other Capniidae . Width of femora fourth to fifth of their length, hind femur reach about the end of Tg 5. Tibiae as long as the femora but less than half the width. Tarsi as wide as tibiae, length of tarsi about half of the corresponding tibia; second tarsomere much shorter than the others, basitarsus shorter than metatarsus on foreleg while as long as on hind leg. Claws symmetrical, smooth and gradually curved, arolium relatively small.

Wings: Macropterous wings about as long as the body, covering more than half of the cerci; wing venation show little variability ( Figs. 56–57 ). Forewing: C simple, narrowing gradually from R4+5 towards Cu1; Sc parallel to C, there are two crossveins between C and Sc besides h, Sc curves abruptly beyond the last crossvein and join R around r; h is closest to arc than to wing base; R1 is distinctly curved between its branching with Rs and r, gives one leaning crossvein towards C shortly after r or at least before halfway to C; R2+3 and R4+5 branching at r or with a short crossvein between r and r-m; M branch out to M1+2 and M3+4 with r-m before r; there is only one crossvein between M and Cu1 besides arc and m-cu, this crossvein continues in cu between Cu1 and Cu2; Cu2 ends before half of the wing length, very narrow or interrupted before arc; usually a thin cu-a presents between Cu2 and A1 around a; A1 ends around the R end of arc, distinctly curved and usually thickened beyond a; A2 reaches only as far as h is positioned. Hindwing: C, Sc and their crossveins similar to the forewing; R1 is straight, not branching with R, but starts separately from arc, gives one leaning crossvein towards C shortly after r or at least before halfway to C; R2+3 and R4+5 branching with a crossvein between r and r-m; M branch out to M1+2 and M3+4 after r-m and m-cu; M thin before branching and there is no crossvein between M and Cu1 between arc and m-cu, but sometimes there is one between M3+4 and Cu1, and cu always present; Cu2 parallel to A 1 in its posterior two thirds and ends beyond the position of r, but very narrow or interrupted before arc; usually a thin cu-a present between Cu2 and A1 around a; anal field large, the fold of the wing extending between the parallel Cu2 and A1; A1 straight beyond a, runs close to Cu2; A2 straight, ends around the position of cu, the nearly longitudinally directed a sometimes not reach A2 but terminates backcurved to the wing base; A3 as long as the distance of arc and wing base, extending perpendicular to the longitudinal veins. In case of micropterous or brachypterous males, the wing length is usually constant in some species but may vary intraspecifically ( Figs. 58–61 ). On these winglets, R is the thickest vein and the wing is folded with about 60° around this vein; arc is distinct on the forewing and sets as far as on the macropterous wings, but usually obscured on the hindwing; single M and Cu run to the wing tip, sometimes also with a few crossveins on the forewing; anal veins of the forewing similar in length to the macropterous ones, usually a normal a and the curve of A1 also conspicuous; anal field of the hindwing proportional enlarged but the veins are vestigial.

Male abdomen: Tg 1–8 entire and unmodified with sinuous antecosta that is entire on Tg 2–9, entire or medially interrupted on Tg 1; transverse row of four spots present on Tg 1–9. Tg 9 with a raised posteromedial process of different shape characteriztic to species; its sclerotization not thicker than the remainder of the tergite, and its caudal face is membranous with specific shape of lateral sclerotization ( Figs. 148–164 ). Tg 10 subdivided with the entire antecosta forming a thick, rectangular U-shaped process in the medial 1/5 of the segment’s width ( Fig. 1 ). Epiproct consists of a large, anteriorly rounded and raised B-scl that is indented medio-apically and divided from both the Ep-scl and the two Lb-scl ( Fig. 1 ); paired Lb-scl small and triangular with sinuous dorsal margin, divided from both the Ep-scl and the B-scl ( Fig. 2 ). Ep-scl long and curved, caudal setae lacking but spines of specific shape and distribution present on the apex; basally wide but abruptly tapering in the basal third; apical two thirds of the sclerite with the same width or medially swollen; dorsally divided in its full length, ventrally connected at the base and the tip, while laterally divided in the apex – concrete shape is characteriztic for the species ( Figs. 106–119 , 127–147 ); Ll lacking; I-scl is a long, dorsally open tube that is curved like the Ep-scl ( Figs. 1–2 ); Ec present in the apical third, between the upper corns of the laterally divided part of Ep-scl ( Figs. 1–2 , 8–10 ).

St 1 entire and unmodified but smaller than the additional sterna, with rounded corners. St 2–8 consist of a large, rectangular posterior and two small anterior sclerites that are declining in size and fused with the posterior sclerite towards the apical segments; a pair of lateral spots present on St 1–8. St 9 reduced to a well sclerotized arch connecting the ventro-basal part of Tg 9; in its medial part, the arch bear a vesicle of variable size from one third of segment’s width to vestigial ( Figs. 83 , 99 ). Sg separated from all other segments, rounded with a more or less triangular shape, apical part with a distinct, incised or rounded tip. Pp wide and short, with rounded apex that is also short and wide. Fp long and narrow, its apical tube curved upward; Rp medium sized, elongated and rounded, divided from Fp ( Fig. 23 ). Cerci longer than the abdomen, with 15 or more segments; segments cylindrical, the basal ones slightly club shaped or pearl-like, gradually elongated towards the apex.

Female abdomen: Tg 1–8 divided into two small, rectangular lateral sclerites; small, hardly sclerotized median plates may also present on Tg 2–6. Tg 9–10 entire and unmodified, Tg 10 posteriorly rounded. St 1 entire and unmodified but smaller than the additional sterna, with rounded corners. St 2–7 consist of a large, rectangular posterior and two small anterior sclerites that are declining in size and fused with the posterior sclerite towards the apical segments. Sg large, covers most of the segment 8 but its posterior end equal to the segment’s end, or only very slightly overhanging; the plate is rectangular with slightly rounded or with broad but very narrow indented posterior margin; sclerotization entire without keel, two small antero-lateral nook usually present ( Figs. 62, 64–65 ). Color of the plate is entirely brown, sometimes a bit paler medially; two small lateral sclerites fused with posteriolateral edges of the Sg. Vaginal complex with broad genital opening, membranous genital cavity reach back to segment 7 where it branching into the oviducts; inner sclerites lacking ( Fig. 63 ). St 9 entire, rectangular; Pp large, having short, rounded tips; epiproct simple, membranous. Cerci as long as or shorter than the abdomen, with 15 or more segments; segments cylindrical, the basal segments slightly clubbed, gradually elongated towards the apex.

Mature larva: Body relatively stout, body length 5.5–12.5 mm. General color pale yellowish brown with very faint pattern; head, dorsal thoracal sclerites besides wing pads, and abdominal terga usually darker. Teneral coloration apparent below cuticle on pharates, together with dark wing pads. Setation long, but inconspicious. Head shape typical of Capniidae , eyes smaller than the area delimited by the three ocelli ( Fig. 66 ). Ocelli and ecdysial suture are prominent, tentorial callosities less pronounced, M-line and occipital rugosities are hard to observe. Mentum small, typical of the Capniidae ; glossae long and apically rounded, paraglossae weakly subdivided, labial palpi short with second palpomere the longest. Maxillae and other mouthparts are of general Capniidae shape and structure, maxillary palpi moderately long with the third palpomere the longest ( Fig. 67 ). Antennae long, with 50 or more antennomeres. Antennomeres cylindrical, the first ten beyond pedicel very short, then gradually elongated towards the apex where antennomeres are about 3× longer than wide. Pn elongated trapezoidal with rounded corners, smooth; rear edge of meso- and metathorax between wing pads rounded, the paired sclerites widely separated from wing pads. Shape of wing pads are typical of Capniidae in macropterous specimens, the mesothoracic pair reach the anterior end of Tg 1, metathoracic pair reach the anterior end of Tg 3; micropterous wing pads always still distinct and separated from the paired sclerites. Lateral thoracic sclerites similar to adults but Epm yet reduced and Aes dorsally not yet divided into Ab, Pb and Al. Ventral thoracic sclerites mostly obscure but Etn, Tn and Fs distinct. PFs longer than of the adult, more elliptical than transverse; MeFs triangular, anterior width half of the intercoxal distance, MeFsa indistinct while MeFsp distinct (forming a Y-ridge with the MeFs); MFs laterally indented ( Figs. 68–69 ). Legs typical of Capniidae , width of femora ⅓ of their length, hind femur reach about the midline of Tg 5. Tibiae as long as the femora but ~½ as wide. Tarsi as wide as tibiae, length of tarsi <½ of the corresponding tibia; basitarsus slightly longer than the second tarsomere on the foreleg, about 2× longer on hind leg but <½ half of the length of metatarsus. Claws symmetrical, smooth and with a basal swelling, regularly curved, arolium vestigial. Abdomen relatively stout, segments 1–9 divided by pleura, integument light and matt. Tg 10 rounded in females, projecting in males; length of the latter equal or shorter than length of Tg 8 and 9, tip blunt, or only slightly raised upwards ( Figs. 74–75 ). Pp wide and short, tip blunt. Cerci usually longer than the abdomen, with 30 or more segments; segments cylindrical, the basal ones short but gradually elongated towards the apex where segments are very slender; the width of segments 11–13 is half to third of their length ( Figs. 76–77 ).

Setation: Head with not so dense, moderately long and thick setae and hairs; eye bear small elongated setae between the ocelli; mouthparts with scarce setation besides the sensilla and apical spines but labrum with a pronounced apical tuft of setae rows ( Figs. 66–67 ). Antennal segments with apical whorl of short setae and hairs; campaniform sensillae occur on their distal ½. Dorsal surface of thorax with similar setation like the head, Pr lacks marginal row or longer marginal setae; setae placed in lines on wing pads. Ventral surface of thorax with scarce, thin and short hairs and setae ( Figs. 68–69 ). Legs with dense setation. All femora bear long, acute setae and thin hairs, and armed with a row of long, erect setae on both dorsal and ventral edges; length of these setae are less than the half of the femur’s width ( Figs. 72–73 ). A bald median line is conspicuous on the dorsal surface of all femora. Tibiae and tarsi covered with long hair-like setae, strong setae present only on the ventral edges of the tibiae; tibiae lack apical spines or the spines are vestigial, scarce dorsal fringe of swimming hairs usually present. Tergal segments with moderately dense, uniform long and blunt setae; apical row lacking, length of the setae are fourth to fifth of segment’s length ( Fig. 70 ). Projection of male Tg 10 having a bare medial line and bare at the apex ( Figs. 74–75 ). Ventral surface of abdomen with fewer but similar setae like tergites; setae dense on the Pp ( Fig. 71 ). Cercal segments with apical row of acute, long setae, short setae and thin hairs; campaniform sensillae occur on the whole surface ( Figs. 76–77 ). Longest setae are ½ to full length of the segment on segments 11–13, as long as or slightly longer than the segment on the apical ones.

Drumming: A comprehensive characterization of the drumming signals used in the genus cannot be given until the drumming signals of Z. sevanica , Z. tuberculata , Z. turkestanica turkestanica , Z. turkestanica brevicula are unknown. The presently known male drumming calls are monophasic beat-groups with gradually decreasing or increasing inter-beat intervals. Solitary males produce single calls sporadically or 1–4 calls in a sequence. The beat repetition patterns ( Figs. 169–184 ) of male calls are species-specific. Male female drumming signal exchanges are male call—female answer or male call—female answer- male response duets ( Figs. 185–189 ).

Genetics: Sequencing coxI of 64 Zwicknia specimens yielded 29 distinct haplotypes with 123 informative characters. Same topology was found with both MP (L = 441, CI = 83, and IR = 93) and ML (ln L =—2470.19612; Fig. 192 ) approaches. Overall divergence among new haplotypes was 6.5 %. Our phylogeny reflected the spatial arrangement of haplotype clusters in the landscape with each newly defined species forming a monophyletic haploclade. In GenBank data analyzed under cox1, the closest related clade was Allocapnia sp. with 89% sequence similarities, and was used as an outgroup.

Affinities. In the West Palaearctic, the genus is distinguishable from other Capniidae in features discussed in the diagnosis of the male and female terminalia. Males of Zwicknia are apparently more closely related to the West Nearctic Bolshecapnia , but differ with having long, undivided tube-like I-scl instead of divided, and the apical part of the Pp short and wide instead of long and narrow. Females of Bolshecapnia have narrower Sg that is usually pointed and overhanging St 8, instead of entire, rectangular Sg with length equal to St 8. Zwicknia is also closely related to the montane Asian Capnia s.l. cordata species group, but the males differ by their lateroapically divided Ep-scl instead of entire, presence of Ec instead of its absence, and the apical part of the Pp short and wide instead of long and narrow. Females of the C. s.l. cordata species group have narrow, dark Sg instead of entire and similar in color to sterna. Larvae are difficult to separate from other Capniidae and an adequate diagnosis cannot be given here. Since no specific characters useful for separation were found, larvae of the species included are not described, only a general description is given. However, matured male larvae usually can de identified on the basis of the epiproct transparent through larval skin. Drumming signals are known only in a small proportion of species in Capniidae . Therefore it would be too early to make a detailed examination of the affinities of Zwicknia species regarding their drumming signals. Mitochondrial DNA showed that Zwicknia species have a range of 19%–22% of overall divergence with respect of Capnia s.l. vidua and a decrease of glutamine of 68% in nucleotide composition, compared with others species of Capniidae .

Distribution and ecology. The genus has a Palaearctic distribution. While common in most parts of Europe, it is absent from North Africa, the Mediterranean Isles, and the Far North ( Iceland, northern Scandinavia). It is reported from the Levant only from Lebanon, but widespread in the northern part of Anatolia, the Caucasus, and Transcaucasia. At least two species enter the East Palaearctic as far as Middle Asia ( Fig 196 ). Adults occur from the second half of January to June. The peak of emergence is in March in most areas. European species inhabit usually slow, medium-sized forest streams of hilly and submontane regions, but they were reported also from small temporary brooks and even high montane streams. Asian species were found generally in later periods and higher elevations up to 2,400 m.

Etymology. The genus is dedicated to Prof. Peter Zwick, Schlitz, Germany, in recognition of his major contributions and establishing leadership in many fields of study of the Plecoptera . The name composed with the ending “nia” corresponding to Capnia ; gender feminine.

Morphological key to adult males of Zwicknia

1 Ep-scl medially swollen ( Figs. 110 , 142–143 ). Process of Tg 9 very wide and distinctly bicornuated ( Fig. 160 ).... Z. kovacsi

- Ep-scl not swollen medially. Process of Tg 9 narrower and without distinct corns................................... 2

2 Tip of Ep-scl upcurved in lateral view ( Figs. 106–107 , 132, 134, 136, 138–141 ). Ep-scl wide and blunt in dorsal view ( Figs. 114–115, 117 ). Spines present only on the membranous part of the epiproct apex ( Figs. 106–107 )...................... 3

- Tip of Ep-scl straight or slightly curved down in lateral view. Ep-scl narrow and more or less pointed in dorsal view. Spines present also on the apical part of Ep-scl....................................................................4

3 Process of Tg 9 low and caudally projecting ( Figs. 92 , 157–159 )...................................... Z. rupprechti

– Process of Tg 9 high and perpendicularly elevated ( Figs. 84 , 148–151 )................................... .. Z. bifrons

4 Tip of Ep-scl acute in lateral view ( Figs. 108–109 , 127–131 ). Process of Tg 9 very high, rectangular and with indenting sides caudally ( Figs. 152–156 )........................................................................... Z. acuta

- Tip of Ep-scl blunt in lateral view. Process of Tg 9 lower, rounded and sides not indenting caudally.................... 5

5 Tip of Ep-scl straight in lateral view ( Figs. 111 , 144–145 ). Process of Tg 9 slightly wider than the epiproct ( Fig. 94 ).................................................................................................... Z. sevanica

- Tip of Ep-scl slightly downcurved in lateral view. Process of Tg 9 obviously wider than the epiproct.................... 6

6 Process of Tg 9 perpendicularly elevated ( Fig. 102 ). Ep-scl comparatively more slender ( Fig. 102, 104–105 , 147 )................................................................................................... Z. turkestanica

- Process of Tg 9 slightly folded backwards ( Fig. 100 ). Ep-scl comparatively more robust ( Figs. 98, 100–101 , 146 ).................................................................................................... Z. tuberculata

Figures

FIGURE 49. Dorsal habitus of male holotype Zwicknia kovacsi Murányi & Gamboa, sp. n.; Romania, Maramureş County, Rodna Mts., Borşa-Staţiunea Borşa—scale 1 mm.

FIGURE 50. Ventral habitus of male holotype Zwicknia kovacsi Murányi & Gamboa, sp. n.; Romania, Maramureş County, Rodna Mts., Borşa-Staţiunea Borşa—scale 1 mm.

FIGURE 51. Lateral habitus of male holotype Zwicknia kovacsi Murányi & Gamboa, sp. n.; Romania, Maramureş County, Rodna Mts., Borşa-Staţiunea Borşa—scale 1 mm.

FIGURE 52. Dorsal pattern of teneral male Zwicknia bifrons (Newman, 1838); Hungary, Pest County, Pilis Mts., Szentendre—scale 1 mm.

FIGURES 53–55. Thoracal sclerites of male Zwicknia kovacsi Murányi & Gamboa, sp. n.—53: ventral view; 54: lateral view; 55: dorsal view (P: prothoracic; Me: mesothoracic; M: metathoracic—Ab: anterior basalare; Aes: anepisternum; Al: alifer; Bs: basisternum; Cx: coxa; Epm: epimeron; Eps: episternum; Etn: eutrochantin; Fs: furcasternum; Fsa: furcasternal arm; Fsp: furcasternal pit; Kes: katepisternum; Lcs: lateral cervical sclerite; Pb: posterior basalare; Pfs: postfurcasternum; Pn: pronotum; Po: postcoxal bridge; Pr: precoxal bridge; Pra: prealar bridge; Prs: presternum; Psc: prescutum; Pscl: postscutellum; Sct: scutum; Scl: scutellum; Ss: spinasternum; St1: Sternite 1; Tg1: Tergite 1; Tgl: tegula; Tn: trochantin; Tr: trochanter—wings and subalar sclerites omitted)—not to scale.

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 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 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 86–89. Male terminalia of Zwicknia kovacsi Murányi & Gamboa, sp. n.; paratype, Romania, Maramureş County, Rodna Mts., Borşa-Staţiunea Borşa—86: dorsal view; 87: ventral view; 88: lateral view; 89: caudal view—scale 1 mm.

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.

FIGURES 94–97. Male terminalia of Zwicknia sevanica (Zhiltzova, 1964); Turkey, Ankara Province, Akyarma Pass—94:

FIGURES 98–101. Male terminalia of Zwicknia tuberculata (Zhiltzova, 1964); Kazakhstan, Jambyl Province, Karatau Mts., Bajaldyrtal—98: dorsal view; 99: ventral view; 100: lateral view; 101: caudal view—scale 1 mm.

FIGURES 102–105. Male terminalia of Zwicknia turkestanica turkestanica (Kimmins, 1950); Kazakhstan, East Kazakhstan Province, Tarbagatai Mts., Ayagos—102: dorsal view; 103: ventral view; 104: lateral view; 105: 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 110–113. Lateral view of male epiproct of Zwicknia Murányi, gen. n.—110: Z. kovacsi Murányi & Gamboa, sp. n.; 111: Z. sevanica (Zhiltzova, 1964); 112: Z. tuberculata (Zhiltzova, 1964); 113: Z. turkestanica turkestanica (Kimmins, 1950)—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 124–126. Male terminalia of Zwicknia Murányi, gen. n., dorso-caudal view—124: Z. sevanica (Zhiltzova, 1964); 125: Z. tuberculata (Zhiltzova, 1964); 126: Z. turkestanica turkestanica (Kimmins, 1950)—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.

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 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.

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.