Isophya kraussii moldavica Iorgu and Heller

Iorgu, Ionuţ Ştefan & Heller, Klaus-Gerhard, 2013, The bush-cricket Isophya kraussii (Orthoptera: Phaneropteridae): bioacoustics, distribution and description of a new subspecies from Romania, Zootaxa 3640 (2), pp. 258-269 : 261-266

Isophya kraussii moldavica Iorgu and Heller , subsp. nov.

( Figs. 1 c, d, e, f; 2 b, d, f, h, j, l, n, p, r; 3 c, d, e; 4 e, f, g, h, m, n, o, p, r; 5)

Type material. Holotype, male, Romania: Suceava, Adâncata, 47°42'17.87"N, 26°18'54.88"E, 400 m, 2007.07.13. Paratypes: 1 ♂, Romania: Suceava, Călineşti, 47°42'02.86"N, 26°18'35.13"E, 410 m, 2007.07.14; 3 ♂ 2 ♀, Romania: Suceava, Adâncata, 47°42'02.86"N, 26°18'35.13"E, 420 m, 2008.07.24; 5 ♂ 2 ♀, Romania: Suceava, Adâncata, 47°42'33.57"N, 26°17'32.21"E, 450 m, 2011.06.24; 2 ♂, Romania: Suceava, Vârfu Dealului, 47°36'56.39"N, 25°56'00.68"E, 500 m, 2011.06.24; 3 ♂ 1 ♀, Romania: Suceava, Arbore, 47°42'53.12"N, 25°52'33.38"E, 430 m, 2011.06.24; 5 ♂ 3 ♀, Romania: Suceava, Putna, 47°53'55.52"N, 25°36'48.09"E, 520 m, 2011.06.24; 7 ♂ 3 ♀, Romania: Suceava, Adâncata, 47°42'25.50"N, 26°17'26.63"E, 450 m, 2012.06.25; 1 ♂ 1 ♀, Romania: Suceava, Botoşana, 47°39'44.08"N, 25°55'33.18"E, 400 m, 2012.06.25; 2 ♂, Romania: Botoşani, Hilişeu–Crişan, 48°00'59.36"N, 26°15'25.31"E, 210 m, 2012.06.25; 3 ♂ 2 ♀, Romania: Botoşani, Lozna, 47°56'48.14"N, 26°18'21.40"E, 300 m, 2012.06.25; 2 ♂ 1 ♀, Romania: Botoşani, Gorovei, 47°52'49.38"N, 26°21'07.64"E, 350 m, 2012.06.25, leg. I. Ş. Iorgu, all in coll. “Grigore Antipa” National Museum of Natural History, Bucharest.

Audio recordings. 3 ♂, Adâncata, 2009.07.0 4, temperature 26°C; 2 ♂, Adâncata, 2010.07.0 3, temperature 20°C; 5 ♂, Adâncata, 2011.06.24, temperature 24°C; 2 ♂, Vârfu Dealului, 2011.06.25, temperature 22°C; 4 ♂, Arbore, 2011.06.25, temperature 17°C; 4 ♂, Putna, 2011.06.26, temperature 22°C; 10 ♂, Adâncata, 2012.06.25, temperature 19°C (IIŞ).

Etymology. The name derives from the historical province of Moldavia, region where the bush–crickets were found.

Diagnosis. Male song consists of long sequences of syllables, each syllable formed of 30–58 impulses and lasting for 89–200 ms. After 60–113 ms, the syllable is usually followed by 1 (extremely rare 2–3) after–clicks. In male, fastigium verticis narrower than scapus, tegmina widened, shorter than pronotum, the right margin of left tegmen forms an angle of about 90° at distal end of Cu2. Stridulatory file with 195–229 pegs. Cerci slightly curved in distal 1/4. Ovipositor upcurved, 10.9–12.1 mm long.

Description. Male. Head with fastigium verticis slightly tapering frontward, narrower than scapus, dorsal sulcus present ( Fig. 2 b). Pronotum saddle–like, disc constricted in the transverse sulcus area, with lateral carinae widen in metazona. Dorsal margin of paranota slightly concave, anterior edge straight, posterior and ventral margins moderately convex ( Figs. 2 b, d). Tegmina shorter than pronotum, angle between cubital veins is about 70°, length of Cu2 vein about 4/5–5/6 posterior border of pronotum, mirror large and quadrangular. At distal end of Cu2, the right margin of left tegmen forms an angle of about 90° ( Fig. 2 b). Stridulatory file length about 2.8–3.1 mm, counting 195–229 teeth ( Fig. 3 c, d). Cercus hairy, gradually narrowing towards tip, curved in apical 1/4. Terminal denticle triangular, positioned in middle of cercus apex; epiproct about 1.7 times as wide as long ( Fig. 2 f). Subgenital plate elongated and narrowed apically, with triangular apical shaped incision ( Fig. 2 h). Hind femur about 4.4–4.8 times pronotum length, without ventral spines ( Table 1 ). Coloration green, with fine dark green spots. In rare cases, males have two dorsal bilateral stripes from pronotum to end of abdomen, dark red or orange colored. Antennae greenish or reddish–brown, with light green scapus. Femora, tibiae and tarsi green, brownish or reddish. A white band begins behind the compound eye and ends at posterolateral area of tegmina. A brown band is present above the white one in metanotum. Tegmina brown, rarely dark green, apical area green and costal margin yellowish–white. Cerci brown or reddish–brown, rarely green.

Female. Fastigium narrower than scapus ( Fig. 2 j). Pronotum with lateral carinae straight, disc marginally enlarged in posterior part and paranota slightly as in males, with ventral margin straight ( Figs. 2 j, l). Tegmina about 1/3 pronotum length, reaching the posterior margin of first abdominal tergite. Right tegmen with stridulatory bristles located on and near cubital veins ( Fig. 3 e). Cercus short, hairy, tapering; epiproct about 1.4 times as wide as long ( Fig. 2 n). Subgenital plate rounded, triangular, about 2.7 times as wide as long ( Fig. 2 p). Ovipositor 2.4–2.7 times pronotum length, upcurved, upper margin with 8–9 denticles and lower margin with 7–8 denticles ( Fig. 2 r). Hind femur 3.6–3.9 times pronotum length, without ventral spines ( Table 1 ). Coloration of body as in males, ovipositor green.

Bioacoustics. Males produce their song usually at dusk and during the night, a long sequence of simple syllables. A syllable is formed of a compact series of 30–58 impulses, impulse interval 2–4 ms, and lasts for about 89–200 ms (n=200 syllables from 30 males). Usually a syllable is followed after 60–113 ms by an after–click, rarely 2–3. The following syllable begins 69–292 ms later ( Figs. 4 e–h, m–p; table 2). Both parts (compact series of impulses and after-click) are the result of one closing stroke of tegmina. So far, we did not record any female acoustic response to male song. Sound frequency ranges from 15–20 kHz up to 40–45 kHz ( Fig. 4 r).

Comparative note. Isophya kraussii moldavica ssp. n. differs from Isophya kraussii kraussii mainly in acoustics and less in morphology. The song of males from the Moldavian populations consists of shorter syllables, formed of 30–58 impulses (mean±SD: 48.24±9.42) and lasting for about 89–200 ms (mean±SD: 114.96±34.87), while males of the nominal subspecies produce longer syllables, formed of 80–125 impulses (mean±SD: 96.97±22.09) and lasting for about 250–443 ms (mean±SD: 313.97±39.5), sometimes up to 600 ms (Roesti & Keist 2009). After–clicks are produced in a similar time window in the 2 subspecies, after 60–113 (mean±SD: 89.26±14.53) in Moldavian individuals and after 41–183 (mean±SD: 92.27±33.78) in Central European specimens. Male stridulatory file is shorter and contains 195–229 teeth in I. kraussii moldavica ssp. n., compared with 260– 305 teeth in I. kraussii kraussii . Some other minor morphological variations were noticed in the two subspecies: male wing with longer and triangular posterior lobes in I. kraussii kraussii and shorter and rounded lobes in I.

kraussii moldavica ssp. n., longer pronotum and wings in males of I. kraussii kraussii and stubbier cercus in males of I. kraussii moldavica ssp. n. Females of the new subspecies have somewhat longer ovipositor: 10.9–12.1 mm, compared with 9.32–11.88 mm in nominal subspecies.

Distribution. Isophya kraussii kraussii is known to occur in Central and Southern Germany, Czech Republic, Slovakia, Southern Poland, Eastern Austria, Eastern Slovenia, Northern Croatia, Western and Northern Hungary. According to Strätz & Königsdorfer (2003) there are a few isolated localities in Bavaria south of the river Danube, where the species is mentioned to have been recorded by three authors before 1950. However, in one of the papers (Kühlhorn 1953), the species (and genus) is not mentioned at all (possibly a mix–up with Tetrix (as Acrydium ) kraussi which is mentioned exactly and only for these localities listed by Strätz & Königsdorfer). In another locality (Höllriegelskreuth) mentioned by Knoerzer (1942) (and based on Knoerzer by Fischer 1950), besides two nymphs, the only adult specimen was beaten from a tree, a quite atypical place for an Isophya . Here possibly an unusually coloured female of Barbitistes was involved. In the same paper, Knoerzer (1942, p. 629) mentions indirectly a previous exchange between I. kraussii (as I. pyrenaea ) and B. serricauda . Therefore these southern Bavarian records are mostly wrong, with one doubtful locality, and they are no shown in the map ( Fig. 5 ). In Czech Republic and Slovakia, Isophya kraussii kraussii is locally common across entire territory, within altitude range 100–1800 m (Kočárek et al. 2005, Holusa et al. 2013, A. Krištín and P. Kočárek pers. comm. 2012).

Isophya kraussii moldavica ssp. n. is known so far only from the hilly meadows in NE Romania, but a record of I. pyrenaea from Central Moldavia by Bey-Bienko (1954), based on two females with unusually long ovipositor, may indicate this subspecies. For the old records of I. pyrenaea from western Ukraine (Łomnicki 1878, 1879 fide Bey-Bienko 1954; Ramme 1951) it is impossible to decide to which subspecies they may belong, if they refer to I. kraussii at all. In 1970, Kis and Vasiliu discussed that most of the early mentions of Isophya pyrenaea in Romania are erroneous, e.g. specimens from “Siebenbürgen Karpaten” (Transylvanian Carpathians) in coll. A. Müller, Brukenthal National Museum, Sibiu. The authors indicate that Isophya pyrenaea is found in Romania only in N Moldavia and the individuals from other regions most probably belong to I. brevipennis Brunner von Wattenwyl (= I. camptoxypha ) or I. modestior Brunner von Wattenwyl. Specimens mentioned by Ramme (1951) from “Königstein bei Braşov” (Piatra Craiului Mountains) may belong to Isophya harzi Kis (Iorgu et al. 2012) .

Ecology. In Germany, Isophya kraussii kraussii is found in a variety of habitats. Common to most of them is a vegetation structure dominated by high grasses or tall herbs and shrubs (Strätz & Königsdorfer 2003). In southern Germany (Bavaria) the species inhabits altitudes between 300 and 900 m (Strätz & Königsdorfer 2003). In Czech Republic and Slovakia, this bush–cricket is considered a submountain and mountain species (Krištín & Kaňuch 2007), occuring in shrubs, taller herbs (Kočárek et al. 2005, Holuša 2006) and steppic xerothermic forest ecotone (Krištín et al. 2011). In Hungary, Isophya kraussii kraussii prefers meso–xerophytic grasslands and forest clearings, ecotone, forest steppe and rocky shrublands with mainly low–shrub species (Rácz 1998, Rácz 2001, Bauer & Kenyeres 2006). In the Bieszczady Mountains from Poland, it was reported from valley, subalpine meadows and clearings (Theuerkauf et al. 2005). Mesophytic ecotone and meadow vegetation with Centaurea , Urtica , Rubus , Rumex , Senecio , Galium , Stachys , Lamium etc. represent the habitat of Isophya kraussii moldavica ssp. n. in Romania. A rich Orthoptera fauna occurs in these types of biotope: Phaneroptera falcata (Poda) , Leptophyes albovittata (Kollar) , Tettigonia viridissima (Linnaeus) , Decticus verrucivorus (Linnaeus) , Metrioptera roeselii (Hagenbach) , Metrioptera bicolor (Philippi) , Pholidoptera griseoaptera (De Geer) , Gryllus campestris (Linnaeus) , Euthystira brachyptera (Ocskay) , Chrysochraon dispar (Germar) , Omocestus rufipes (Zetterstedt) , Stenobothrus lineatus (Panzer) , Chorthippus apricarius (Linnaeus) , Chorthippus biguttulus (Linnaeus) , Chorthippus dorsatus (Zetterstedt) , Chorthippus parallelus (Zetterstedt) etc. The syntopic occurrence of Isophya species remains an interesting subject, I. kraussii kraussii and I. pienensis (Mařan) being found together in Bieszczady Mountains, Poland (Theuerkauf et al. 2005). The occurrence of Isophya kraussii moldavica ssp. n. syntopic with I. pienensis in the Eastern Subcarpathians and with I. zubowskii (Bey–Bienko) in the Moldavian plane W of Prut river are not to be excluded, although we have not found these associations so far.

Figures

FIGURE 2. Morphological details. Isophya kraussii kraussii: a. male head, pronotum and wings (dorsal view); c. male pronotum and wings (lateral view); e. male cerci; g. male subgenital plate; i. female head, pronotum and wings (dorsal view); k. female pronotum and wings (lateral view); m. female cerci; o. female subgenital plate; q. ovipositor; Isophya kraussii moldavica ssp. n.: b. male head, pronotum and wings (dorsal view); d. male pronotum and wings (lateral view); f. male cerci; h. male subgenital plate; j. female head, pronotum and wings (dorsal view); l. female pronotum and wings (lateral view); n. female cerci; p. female subgenital plate; r. ovipositor. Scale unit: 1 mm.

FIGURE 3. Scanning electron microscope photos of stridulatory file: a. Isophya kraussii kraussii male (ch 1907); b. Isophya kraussii kraussii male (chx 214); c. Isophya kraussii moldavica ssp. n. male (Putna); d. Isophya kraussii moldavica ssp. n. male (Adâncata); e. Isophya kraussii moldavica ssp. n. female (Adâncata). Scale 1 mm.

FIGURE 4. Sound analysis in Isophya kraussii kraussii (a – d, i – l, q) and Isophya kraussii moldavica ssp. n. (e – h, m – p, r): a – h, male song oscillogram; i – p, detailed syllable oscillogram; q, r, spectrum (a, i, light blue line, Würzburg, Germany, 23.5 ºC (ISKR 8109); b, j, brown line, Altheim W Neustadt / Aisch, Germany, 21.5 ºC (ISKR 8401); c, k, black line, Nordbayern, Germany, 27 ºC (ISKR 8613); d, l, purple line, Krapina, Ptuj, Croatia, 26 ºC (ISKR 8813); e, m, red line, Adâncata, Romania, 24 ºC; f, n, green line, Arbore, Romania, 17 ºC; g, o, dark blue line, Putna, Romania, 22 ºC; h, p, orange line, Vârfu Dealului, Romania, 22 ºC).

FIGURE 5. Distribution map of Isophya kraussii kraussii (black circles), Isophya kraussii moldavica ssp. n. (white circles) and Isophya pyrenaea (old literature data, most probably = Isophya kraussii, black triangles), based on original and literature data (Bauer & Kenyeres 2006, Bedjanič 2009, Bey-Bienko 1954, Buchweitz 1993, Chládek 1984, Chládek 1986, Chládek 1988, Chládek 1995, Chládek 2001, Čejchan 1955, Čejchan 1959, Čejchan 1986, Čejchan 1987, Čejchan 1988 a, Čejchan 1988 b, Čejchan 1989, Čejchan 1993, Čejchan 1994, Derbuch & Berg 1999, Fikáček 2012, Grzywacz & Warchałowska – Śliwa 2008, Heller 1988, Heller et al. 2004, Holuša 2006, Ingrisch 1991, Kenyeres et al. 2009, Krištín 2001, Krištín & Hrúz 2005, Krištín & Mihál 2000, Krištín & Kaňuch 2007, Krištín & Kaňuch 2013, Krištín et al. 2009, Krištín et al. 2011, Laußmann 1993, Maas et al. 2002, Merkel 1941, Nagy et al. 1998, Nagy et al. 2003, Nagy et al. 2007, Pecsenye et al. 2003, Rácz 1998, Schreiber 1996, Steglich & Müller 2011, Theuerkauf et al. 2005, Warchałowska – Śliwa et al. 2008, Zacher 1917, Zechner & Fachbach 2001, Zuna – Kratky et al. 2009).

FIGURE 1. a. Isophya kraussii kraussii male (Germany, Bayern, Ailsbachtal, 1981.06. 21, KGH); b. I. kraussii kraussii gynandromorph specimen (Germany, Bayern, Münnerstadt, 1982.06. 30, KGH); c. I. kraussii moldavica ssp. n. male (Romania, Hilişeu – Crişan, 2012.06. 25, IIŞ); d. I. kraussii moldavica ssp. n. female (Romania, Hilişeu – Crişan, 2012.06. 25, IIŞ); e. habitat at Adâncata, Romania, 2011.06. 24 (IIŞ); habitat at Arbore, Romania, 2011.06. 24 (IIŞ).

Tables

TABLE 1. Comparative morphological characters in Isophya kraussii subspecies.

			HW (mm)	PL (mm)	PW (mm)	TL (mm)	TW (mm)	BL (mm)	FL (mm)	CL (mm)	S	O (mm)
kraussii kraussii Isophya	male (n=7) female (n=3)	Max. Min. Mean S. D. Max. Min. Mean	3.3 3.13 3.24 0.08 3.72 3.31 3.57	4.18 3.9 4.01 0.12 4.51 4 4.25	4 3.81 3.90 0.09 4.35 3.5 3.94	3.59 3.44 3.5 0.07 1.91 1.35 1.68	3.97 3.75 3.88 0.10 3.23 2.9 3.11	23.9 19.3 21.57 1.39 23.4 19.71 21.7	15.39 14.55 15 0.43 16.4 15.09 15.66	2.39 2.27 2.32 0.06 1.29 1 1.15	274 296 287 8.79 – – –	– – – – 11.88 9.32 10.69
		S. D.	0.23	0.26	0.43	0.30	0.18	1.86	0.67	0.15	–	1.29
. n. ssp moldavica kraussii Isophya	male (n=15) female (n=15)	Max. Min. Mean S. D. Max. Min. Mean S. D.	3.69 3.27 3.49 0.12 4.1 3.85 3.94 0.07	3.92 3.3 3.63 0.19 4.76 4.29 4.49 0.15	3.96 3.85 3.89 0.03 4.25 4.14 4.19 0.03	3.56 3.33 3.43 0.07 1.98 1.5 1.73 0.14	3.72 3.57 3.64 0.05 3.4 3.11 3.23 0.09	25 22.3 23.62 0.70 24.95 23.3 24.15 0.44	17.8 15.9 16.66 0.55 17.55 16.8 17.15 0.22	2.74 2.24 2.42 0.13 1.35 1.19 1.25 0.06	229 195 201.93 8.15 – – – –	– – – – 12.1 10.9 11.59 0.52