Romania, Eastern Carpathian Mountains, Călimani Mountains.

Holotype: male. Original label: “Romania, Suceava County, Călimani Mountains, 12 Apostoli Peak, 47.221233°N, 25.213896°E, alt. 1730 m asl, 2015.07.27, leg. I. Ș. Iorgu”, coll. “Grigore Antipa” National Museum of Natural History, Bucharest, Romania. Paratypes: 1 ♂ 1 ♀, labeled: “Romania, Suceava County, Gura Haitii, 47.226236°N, 25.300690°E, alt. 990 m asl, 2012.06.21, leg. I. Ș. Iorgu”; 2 ♂♂ 2 ♀♀, labeled: “Romania, Suceava County, Călimani Mountains, meadow below Pietrosul Peak, 47.130767°N, 25.191679°E, alt. 1800 m asl, 2012.07.20, leg. I. Ș. Iorgu”; 1 ♂ 1 ♀, labeled: “Romania, Suceava County, Suhard Mountains, Omu Peak, 47.507519°N, 25.090278°E, alt. 1860 m asl, 2013.07.21, leg. I. Ș. Iorgu”; 2 ♂♂ 2 ♀♀, labeled: “Romania, Suceava County, Călimani Mountains, 12 Apostoli Peak, 47.221233°N, 25.213896°E, alt. 1730 m asl, 2015.07.27, leg. I. Ș. Iorgu”; 2 ♀♀, labeled: “Romania, Suceava County, Călimani Mountains, 12 Apostoli Peak, 47.222835°N, 25.212571°E, alt. 1730 m asl, 2016.09.03, leg. I. Ș. Iorgu”, all in coll. “Grigore Antipa” National Museum of Natural History, Bucharest, Romania; 3 ♂♂ 3 ♀♀, labeled: “Romania, Suceava County, Călimani Mountains, 12 Apostoli Peak, 47.221233°N, 25.213896°E, alt. 1730 m asl, 2015.07.27, leg. I. Ș. Iorgu”, coll. Hungarian Natural History Museum, Budapest, Hungary.

Audio recordings: 1 ♂, Gura Haitii, 2012.06.21 (air temperature 25°C); 1 ♂, meadow below Pietrosul Peak, Călimani Mts., 2012.07.20 (22°C); 2 ♂♂ 3 ♀♀, meadow below Pietrosul Peak, Călimani Mts., 2012.07.20 (27°C); 3 ♂♂, Omu Peak, Suhard Mts., 2013.07.21 (25°C); 4 ♂♂ 2 ♀♀, 12 Apostoli Peak, Călimani Mts., 2015.07.27 (24°C).

Figure 1. 

Isophya bucovinensis sp. n.: A female habitus B male habitus C habitat in Călimani Mountains, at 12 Apostoli Peak (1730 m asl) on 27 July 2015.

Isophya bucovinensis sp. n.: 1 ♂ 1 ♀, Romania, Suceava County, Călimani Mountains, 47.208928°N, 25.200489°E, alt. 1640 m asl, 2015.07.27, leg. I. Ș. Iorgu; 2 ♂♂ 3 ♀♀, Romania, Suceava County, Călimani Mountains, 47.202448°N, 25.197817°E, alt. 1620 m asl, 2015.07.28, leg. I. Ș. Iorgu; 1 ♂ 4 ♀♀, Romania, Suceava County, Călimani Mountains, 47.134651°N, 25.178858°E, alt. 1680 m asl, 2016.08.07, leg. I. Ș. Iorgu. Isophya nagyi: 10 ♂♂ 2 ♀♀ (audio recorded at 21.2–24°C), Romania, Suceava County, Călimani Mountains, 47.137°N, 25.256°E, alt. 1340 m asl and 47.141°N, 25.267°E, alt. 1580 m asl, 2011.07.27, leg. K. M. Orci, G. Puskás & G. Szövényi.

Fastigium verticis approximately half as wide as scapus, male elytra short and narrow, slightly shorter than or as long as pronotum, reaching anterior part of second abdominal tergite. Length of cubital vein 3/4 of the width of pronotum posterior margin, forming an obtuse angle of nearly 110° at the right margin of left tegmen. The stridulatory file is 2.2–2.5 mm long and contains 105–130 pegs. Male cerci slender, 2.4–2.7 mm long, gradually and moderately incurved in distal 1/4, tapering in an apical denticle. Ovipositor short, upcurved, 8.3–9.2 mm long. Male calling song consists of a long sequence of two-component syllables (Fig. 3A, B) repeated at a rate of 160–220 per minute (at 22–27°C air temperature). Each syllable lasts for 237–385 ms and is formed of two parts: a longer group of 46–79 impulses (156–286 ms) shortly followed by a higher amplitude group of 2–7 clicks (3–28 ms) (Fig. 3C, D). Females reply right after the male syllable, with a latency of 48–66 ms (Fig. 4C). All these sounds are produced by closing the tegmina.

Morphologically, Isophya bucovinensis sp. n. belongs to the Isophya camptoxypha species group. Within this complex, I. bucovinensis sp. n. and I. nagyi differ from all the other species in the relatively high number of pegs observable in the male stridulatory file. Furthermore, Isophya bucovinensis sp. n. can be easily distinguished from I. nagyi by examining the number of impulse groups within a syllable (three in I. nagyi vs. two in I. bucovinensis sp. n.) and the syllable repetition rate during continuous singing: 60–80 syllables/minute in I. nagyi (21–24°C) and 160–220 in I. bucovinensis sp. n. (22–27°C) (Fig. 4).

Male (Fig. 2A, C, H, J, K; Table 1). Fastigium verticis half as wide as scapus, slightly tapering frontward. Head width 3.1–3.7 mm, 1.3 times the maximum pronotum width. The pronotum is 3.9–5 mm long, with lateral carinae nearly parallel in prozona, broken at traverse sulcus, widen and divergent in metazona. Saddle–shaped from a lateral view, the paranota with concave dorsal margin; anterior and ventral borders straight, posterior edge moderately convex. Tegmina short and narrow, slightly shorter than or as long as pronotum, reaching the second abdominal tergite. Venation reticulate, Cu2 vein swollen, its length 3/4 of pronotum posterior margin. The angle between cubital veins of approximately 70–80°, speculum large and quadrangular. At the distal end of Cu2, the right margin of left tegmen forms an obtuse angle of nearly 110°. Stridulatory file arcuate, 2.2–2.5 mm long, with 105–130 teeth. Hind femur without ventral spines, 3.5–4 times long as pronotum. Epiproct almost twice as wide as long. Cerci covered by fine hairs, slender, 2.4–2.7 mm long, gradually narrowing towards tip, slightly curved in apical fourth. Terminal denticle strong, located in middle of cercus apex. Subgenital plate moderately elongated, narrowed apically with triangular apical incision and more or less acute lobes on its caudal margin.

Figure 2. 

Photographs of the typical eidonomy of Isophya bucovinensis sp. n. male (A, C, H, J, K) and female (B, D, E, F, G, I). A, B head, pronotum and tegmina in dorsal view C D head, pronotum and tegmina in lateral view E ovipositor in lateral view F female subgenital plate G female cerci in dorsal view H male stridulatory file (SEM) I female stridulatory bristles (SEM) J male cercus (SEM) K male cerci in dorsal view. Scale bars 1 mm.

Coloration green, densely punctuated with fine, dark green and brown spots. Some specimens with two dorsal, parallel, bilateral stripes from pronotum to end of abdomen, red, orange, or yellow colored. Antennae greenish, reddish or brownish, compound eyes bicolored: upper part brown or red and lower part green. Disc of pronotum green, bordered by a yellowish or whitish band, from behind the eye to the posterolateral angle of wing, complemented by a reddish brown stripe in metazona. Tegmina brown or dark red, with green margins and greenish or yellowish white costal margin, as continuation of pronotum band. Legs usually green, brownish or reddish. Cerci brown or reddish brown.

Female (Fig. 2B, D, E, F, G, I). Head roughly as in male. Pronotum disc marginally widening in its posterior part, with lateral carinae straight in prozona. Paranota similar to these of male. Tegmina with dense reticulate venation, approximately 1/4–1/3 the length of pronotum, reaching the first abdominal tergite, with more or less rounded margins. Right tegmen with two dorsal fields of stridulatory bristles on cubital veins. Epiproct semicircular. Cercus short, hairy, conical. Subgenital plate rounded, narrow. Ovipositor relatively short, upcurved, 8.3–9.2 mm long, with 8–9 spines on dorsal margin and 7–9 spines on ventral margin, gonangulum ellipsoid.

Body coloration similar to that of male. Tegmina green with brownish inner and whitish or yellowish lateral edges. Ovipositor green with dark brown spines.

(Figs 3, 4; Table 2). Males stridulate in the evening and at night. Typically, the song consists of series of 8–35 syllables; uninterrupted series of more than 200 syllables were also recorded. Syllable repetition rate is 160–220 syllables/minute at 22–27°C. Isophya bucovinensis sp. n. produces a single syllable type, lasting for 237–385 ms (mean ± SD: 311.6 ± 57.85 ms, n = 11 males) and formed of two elements: the main part (“B”) is a longer group of 46–79 impulses (mean ± SD: 60.1 ± 11.54), 156–286 ms (mean ± SD: 215.2 ± 52.7 ms) long, shortly followed by the terminal “C” part, a higher amplitude group of 2–7 impulses (mean ± SD: 4.4 ± 1.64), 3–28 ms (mean ± SD: 16.45 ± 7.39 ms) long. The same impulse interval was measured in both “B” and “C” groups, 2–4 ms. The two parts are separated by a short silent interval of 59–94 ms (mean ± SD: 76.85 ± 10.56 ms). The following syllable begins 76–125 ms (mean ± SD: 94.2 ± 17 ms) later (Fig. 3). All sounds are produced when the male closes the tegmina. Apart from the described typical syllable structure, some males produce syllables without the “C” element at the beginning of syllable series. The calling song’s dominant frequency components are between 10–35 kHz, highest peak at approximately 25 kHz (Fig. 4E).

Figure 3. 

Oscillographic representation of the calling songs (A, B) of two males of Isophya bucovinensis sp. n. and detailed syllables (C, D). A, C Călimani Mountains, 12 Apostoli peak, 1730 m asl (air temperature 24°C) B, D Suhard Mountains, Omu peak, 1860 m asl (air temperature 25°C).

Females find males by phonotaxis and emit simple clicks if the male song is attractive for them. In the resulting male–female acoustic duet, the female replies only after the male syllables (“C”–part), with a latency of 48–66 ms (mean ± SD: 58.7 ± 4.77 ms, n = 200 replies from 5 females) (Fig. 4C).

Figure 4. 

Acoustic signals analysis in two close–related species: A, B oscillographic pattern of male calling song C, D a detailed syllable and the female response (marked with green) E spectrum (function: Hanning window, size: 512). A, C, E red line Isophya bucovinensis sp. n. B, D, E blue line Isophya nagyi.

The specific name is derived from the historical region of Bucovina, “Land of the beech forests”.

The new species populates mesophytic montane meadows in the south–western part of Bucovina, within the altitudinal range of 900–1900 m (Fig. 1C). The insects were collected from broad leaves of various dicotyledonous plants: Veratrum sp., Rumex sp., Rubus sp., Aconitum sp., Vaccinium sp., Urtica sp., Gentiana sp., Hypericum sp. etc. The distribution of this bush-cricket includes highlands from northern and western Călimani Mountains, the northern part of Suhard Mountains (Eastern Carpathians) (Fig. 6) and most likely occurs also in Dorna depression and the central and southern parts of Suhard Mountains.

Isophya bucovinensis sp. n. is known only from Călimani and Suhard Mountains. It could be considered as Endangered B1ab(iii, v)+2ab(iii, v) following IUCN’s Red List categories and criteria version 3.1 (known EOO=234 km² and AOO=28 km²; the constant degradation of species’ habitat is the result of mechanized mowing in June and July, and overgrazing, especially in Suhard highlands) (IUCN 2012).

In this study, we report the discovery and give the description of a new bush-cricket species recognized based on its distinctive calling song. The basic pattern of its song is not unique within the genus Isophya: a series of evenly repeated syllables, each consisting of two elements: a longer impulse series followed by a short impulse group (see e. g. Heller 1988, Chobanov et al. 2013). However, species producing this male song pattern can be found in several different, morphology-based species-groups (Warchałowska-Śliwa et al. 2008, Chobanov et al. 2013). One of these species-groups is the Isophya camptoxypha complex. Six morphologically closely-related species inhabiting the Carpathians were recently placed within this group: I. camptoxypha Fieber, I. ciucasi Iorgu & Iorgu, I. nagyi, I. sicula Orci, Szövényi & Nagy, I. posthumoidalis Bazyluk and Isophya dochia Iorgu (Szövényi et al. 2012, Iorgu 2012). Because of its similar morphology, I. bucovinensis sp. n. also belongs to this group.

Since the species of the Isophya camptoxypha species-group are all very similar to each other regarding the morphology, we believe that a male calling song based identification key will be helpful to identify specimens from populations of unknown species identity.

Identification key for the presently recognized species of I. camptoxypha species group, based on male song characters:

Figure 5. 

Oscillograms showing the typical pattern of male calling songs in each species of the Isophya camptoxypha species-group: A I. posthumoidalis (25°C, Bieszczady Mountains, Poland) B I. dochia (24.8°C, Gurghiu Mountains, Romania) C I. camptoxypha (25°C, Bieszczady Mountains, Poland) D I. sicula (24.8°C, Harghita-Ciceu Mountains, Romania) E I. ciucasi (25°C, Ciucaș Mountains, Romania) F I. nagyi (22.5°C, Călimani Mountains, Romania) G I. bucovinensis sp. n. (25°C, Suhard Mountains, Romania). Time scale (shown at the bottom) is the same for all oscillograms in the figure. The arrows indicate the characteristic interval separating syllable groups.

The duration values given in the above key are based on male songs recorded at an ambient air temperature between 20–25°C. To identify specimens, a song recording should be made in that range of air temperature. The key is most reliable when mean values based on at least 5–10 measurements per specimen are used in all quantitative characters.

The easiest way to identify male specimens of Isophya bucovinensis sp. n. is to examine both the male stridulatory file and the pattern of male calling song. Within the I. camptoxypha species group, only two species have male stridulatory files containing more than 100 stridulatory pegs: I. nagyi and I. bucovinensis sp. n. Interestingly, except for the different number of syllabic elements and syllable repetition rates, the syllables produced by males of Isophya bucovinensis sp. n. and I. nagyi show remarkable similarities in oscillographic structure (Fig. 4). The longest impulse series of their syllables (element “B”) are almost identical (156–286 ms, 46–79 impulses in I. bucovinensis sp. n. and 129–217 ms, 38–79 impulses in I. nagyi), and also the high amplitude, short impulse group (element “C”) after the long impulse series is rather similar in the two species (3–28 ms, 2–7 impulses in I. bucovinensis sp. n. and 1–6 ms, 1–4 impulses in I. nagyi). Because of these similarities in song and due to the high level of morphological similarity, we hypothesize that the two species are likely to be a pair of young sister species. This premise should be examined during subsequent studies relying on molecular genetic analysis. Spectral properties of the male calling songs are similar in these two species, most likely due to the comparable dimensions of male stridulatory apparatus (Fig. 4E).

In Isophya bucovinensis sp. n., the delay of female response is much shorter (48–66 ms) than that in I. nagyi (71–127 ms) (Fig. 4C, D). Since female reply delay proved to be a critical parameter for males to recognize female response as a conspecific signal in a number of phaneropterine bush-crickets (Heller & von Helversen, 1986), the detected difference in response delay is likely to make the responses of I. bucovinensis sp. n. females ineffective for communicating with I. nagyi males and vice versa. Also, another impediment for heterospecific communication in these two species is that the responses of I. nagyi females would overlap with a dense impulse series of the male’s next syllable if she would try to duet with a male I. bucovinensis sp. n.; the detection of the female signal is less likely if the male’s own signal masks it. Therefore, in addition to the clear difference of male songs, the divergence of female response delay also suggests that the two taxa are best treated as specifically distinct. However, several interesting questions arise regarding the male–female communication of these two species. Presently, it is uncertain which feature of the male signal serves as a marker to start the timing of female response. Females of duetting Phaneropterinae bush-crickets usually use a conspicuous motif of the male song as a response timing marker (Heller and von Helversen 1986, Heller 1990, Dobler et al. 1994). Clearly detectable features of the male song in I. bucovinensis sp. n. are the beginning and end of “B” and “C” elements. Among these, we consider the beginning of “C” element to be most likely the female response timing marker (this is why we measured female response delay from that male song feature). However, it is only a presumption and a future laboratory test with playback experiments applying modified male syllables will be needed to answer this question.

The Isophya camptoxypha species-group shows a surprisingly high rate of endemism in the Eastern Carpathians. Five of its recently described species have very small distribution areas (Fig. 6) : I. ciucasi populates montane meadows in the southern part of the Eastern Carpathians: in Ciucaș and Vrancea Mountains, I. dochia is known to occur in Ceahlău, Bistriței and Gurghiu Mountains, I. sicula is distributed in Harghita-Mădăraș and Stânișoarei Mountains, I. nagyi occurs in small areas in the southern part of Călimani volcanic caldera and I. bucovinensis sp. n. is known from the northern and western parts of Călimani, reaching Suhard mountains in north. The ancestors of these species presumably penetrated from the distribution center of this genus in Balkan–Asia Minor along the western part of Southern Carpathians and Apuseni Mountains dispersal routes (Kenyeres et al. 2009), and speciated during their expansion by isolation of the populations in the mountain chains of Eastern Carpathians.

Figure 6. 

Geographic distribution of the Eastern Carpathian endemic Isophya species: red dots – I. bucovinensis sp. n., yellow pentagons – I. ciucasi, green squares – I. dochia, blue diamonds – I. nagyi, black triangles – I. sicula.