Review of the Balkan Isophya (Orthoptera: Phaneropteridae) with particular emphasis on the Isophya modesta group and remarks on the systematics of the genus based on morphological and acoustic data Author Dragan P. Chobanov Author Beata Grzywacz Author Ionuţ Ş. Iorgu Author Battal Cιplak Author Maya B. Ilieva Author Elżbieta Warchałowska-Śliwa text Zootaxa 2013 3658 1 1 81 journal article 39033 10.11646/zootaxa.3658.1.1 0adf09ba-5301-4828-abf6-8dbe89f041ca 1175-5326 246551 C02D1C74-25C0-41DD-B098-62098EB7B62A 2.3. Isophya thracica Karabag 1962 ( Figs 3 , 34 , 59 , 83 , 108 , 134 , 138 , 191 ) Isophya thracica Karabag : Karabag 1962 (sp.n.). Morphological description : Karabag 1962; Harz 1969; Heller 1988. Bioacoustics : Heller 1988. The species has unique combination of characters of the male and female tegmina ( Figs 3 , 34 , 59 ). While males have outstandingly long tegmina with very short CuP similarly to the primitive state in I. straubei group, the female tegmina show advanced shortening and reticulate venation of tegmina. Hind femora lack ventral spines. CuP is weak and among the shortest in Isophya being about 1/2 of the hind edge of pronotum; CuP and CuA are widely separated. Male and female stridulatory apparatus are shown on Fig 134 (A and D). Male stridulatory row has a length of 1.78–2 mm with 58–70 teeth. Male cerci ( Fig. 83 , 134 C) are stout, apically strongly incurved and bear a large, showel-like tooth. The ovipositor ( Fig. 108 ) is short ( 7.5–9 mm ). The song ( Fig. 113 ) consists of short isolated syllables divided into main part and an after-click. FIGURES 56–79. Morphology of female head, pronotum and tegmina of the Balkan Isophya and I. yaraligozi (localities as in Figs 31–55 unless specifically given): 56— I. hospodar ; 57— I. rectipennis ; 58— I. pavelii (paratype of I. rammei Peshev ); 59— I. thracica ; 60— I. bureschi ; 61— I. yaraligozi ; 62— I. tosevski ; 63— I. andreevae (paratype); 64— I. clara ; 65— I. miksici (paratype); 66— I. plevnensis (paratype); 67— I. longicaudata adamovici (paratype, BG: Sliven, 1000 m, NMNHS); 68— I. l. longicaudata ; 69— I. rhodopensis leonorae (BG: “Alibotush, 1500 m”, 13.07.1959, NMNHS); 70— I. rh. rhodopensis (as in Fig. 46 but 17.07.1956); 71— I. rh. petkovi ; 72— I. modestior ; 73— I. dobrogensis ; 74— I. zubowskii ; 75— I. aff. camptoxypha (MK: Bistra Mt., 2.08.2004, CC); 76— I. gulae (paratype, BG: Dolna Topchiya, 10.07.1974, NMNHS); 77— I. obtusa (BG: Vezhen Peak, 15.07.1969, NMNHS); 78— I. amplipennis (TR: “Turkey | 1960 | Sureya Bey”, NHM); 79— I. speciosa (BG: Sakar Mt., 14.06.1952, NMNHS). Scale (if present; black line right of the specimen) = 10 mm. Bioacoustics : The song was studied in specimens from Elmali Village in European Turkey . At a temperature of 17–19°С the song consisted of single syllables that rarely were mixed with groups of two syllables. Syllables were repeated at an interval of 7–15 or, when grouped, at 1.5–3 s. The syllables are composed by main part followed by additional part of a single or up to three after-clicks. The main syllable part lasted 67–86 ms (mean 76±5; n=32) and consisted of 18–25 impulses (mean 21±2; n=30) with impulse period of 3–9 ms. The first impulse has much higher amplitude than the rest in the syllable. The additional syllable part followed at 48–143 ms (mean 82±27; n=31) and was composed by one or up to three impulses; the impulse period if more than one impulse lasted 10–15 ms. The total syllable length was 124–212 ms (mean 174±23; n=31). The song properties presented by Heller (1988) were measured at over 24.5°С and in order to be comparable we preferred to give similar data in the Tabulated key ( Table 1 ). Distribution ( Fig. 191 ) and phenology : Endemic for the southern part of Eastern Thrace (European Turkey ). Its phenology is poorly known; early species emerging in February–April and imaginal moulting from the end of April; imagines disappear in July. Near Elmali Village the species was found within xerothermophyte scrub of Quercus pubescens and Juniperus oxycedrus but possibly inhabits a wide variety of vegetation types . FIGURES 80–104. Male abdominal terminalia of the Balkan Isophya and I. yaraligozi (localities as in Figs 31–55): 80— I. hospodar ; 81— I. rectipennis ; 82— I. pavelii ; 83— I. thracica ; 84— I. bureschi ; 85— I. yaraligozi ; 86— I. tosevski ; 87— I. andreevae ; 88— I. clara ; 89— I. miksici ; 90— I. plevnensis ; 91— I. longicaudata adamovici ; 92— I. l. longicaudata ; 93— I. m. modesta (= intermedia ); 94— I. rhodopensis leonorae ; 95— I. rh. rhodopensis ; 96— I. rh. petkovi ; 97— I. modestior ; 98— I. dobrogensis ; 99— I. zubowskii ; 100— I. aff. camptoxypha ; 101— I. gulae ; 102— I. obtusa ; 103— I. amplipennis ; 104— I. speciosa . Scale (if present) = 2 mm. TABLE 1. Tabulated key for distinguishing the Balkan species of Isophya .

Species/ Character	Width of fastigium / scapus	Width of male tegmina in comparison to width of metazone	Length of male tegmina: length of pronotum	Shape of anal angle of left tegmen	Length of CuP: width of metazone	CuA– CuP position	Position of veins of female tegmina	Number of stridulatory teeth	Shape of male cerci	Length of ovipositor /pronotum	Song structure	Syllables main part (20– 30°C) [ms]	Syllables main + additionalpart (20–30°C) [ms]	Melanism	Colouration of stripes at dorsolateral borders of pronotum	Distribution
I. hospodar	±equal	narrower	1.2–1.5: 1	obtuse	1/2–2/3	widely separated	±parallel	63–78	Fig. 80	2–2.5	groups of phrases	<50	–	absent	no dark stripes	E Balkans
I. rectipennis	±equal	narrower	±equal	obtuse	1/2–2/3	widely separated	±parallel	45–60	Fig. 81	1.6–2	phrases	10–50	–	absent	no dark stripes	SE Romania, E Balkans, NW Anatolia
I. pavelii	±equal	±equal	±equal or slightly longer	obtuse	2/3–3/4	slightly attached	±parallel	97–115	Fig. 82	1.7–2.1	phrases	25–120	–	absent	reddish	SE Balkans, NW Anatolia
I. thracica	1/2–equal	narrower	±1.5: 1	obtuse	±1/2	widely separated	reticulate	58–70	Fig. 83	1.6–<2	groups of syllables	<70	±100	absent	reddish	S European Turkey

I. bureschi	1/2	narrower	±equal or slightly longer	obtuse	1/2–2/3	slightly attached	intermediate	68–75	Fig. 84	>2.3	groups of syllables	30–60	500–>1000	absent	reddish	Locally in C Bulgaria (see text)
I. yaraligozi	±equal	wider	±equal or slightly longer	right	3/4–4/5	distinctly attached	intermediate	±140	Fig. 85	2–2.3	groups of 2 or more syllables	150–300	–	absent	reddish	NW Anatolia – Yaraligoz Mt.
I. tosevski	1/2–2/3	slightly narrower	±equal or slightly longer	obtuse	2/3–3/4	distinctly attached	intermediate	88–110	Fig. 86	2–2.3	isolated syllables	150–250	350–>500	absent	reddish	SW Macedonia, N Greece (Vardar Valley)
I. andreevae I. clara I. miksici	1/2–2/3 1/2–2/3 1/2–2/3	slightly narrower ±equal slightly narrower	±equal or slightly longer ±equal or slightly longer ±equal or slightly longer	obtuse obtuse obtuse	2/3–3/4 1/2–2/3 1/2–2/3	distinctly attached slightly attached distinctly attached	intermediate intermediate intermediate	77–92 58–72 68–75	Fig. 87 Fig. 88 Fig. 89	>2.3>2.5>2.3	groups of syllables groups of syllables groups of syllables	180–450 100–350 100–250	300–>500 – 200–>500	absent absent absent	reddish reddish reddish	SW Bulgaria, N Greece (Strouma Valley) Bosnia and Herzegovina, Montenegro, Serbia NW Bulgaria and neighbouring Serbia
I. plevnensis	1/2–2/3	±equal	±equal or slightly longer	obtuse	2/3–3/4	strongly attached	intermediate	100–130	Fig. 90	>2.4	groups of 2–4 syllables	200–400	>500	absent	reddish	Central N Bulgaria
I. l. adamovici	1/2–2/3	±equal	±equal or slightly longer	obtuse	2/3–3/4	strongly attached	intermediate	145–160	Fig. 91	>2.5	isolated syllables	400–>850	–	absent	reddish	E Stara Planina Mts (E Bulgaria)
I. l. longicaudata	1/2–3/4	±equal	±equal or slightly longer	obtuse	3/4–4/5	strongly attached	intermediate	145–160	Fig. 92	>2.5	isolated syllables	130– 200	>850– 1500	absent	reddish	E and NE Bulgaria and neighbouring Romania
I. modesta	1/2–2/3	±equal	±equal or slightly longer	obtuse	2/3–3/4	distinctly attached	intermediate	124–143	Fig. 93	>2.4	isolated syllables	120–240	5000–>10000	absent	reddish	C and E Europe

…… continued on the next page II. Northern stock or Northern species of Isophya TABLE 1. (Continued)

Species/ Character	Width of fastigium / scapus	Width of male tegmina in comparison to width of metazone	Length of male tegmina: length of pronotum	Shape of anal angle of left tegmen	Length of CuP: width of metazone	CuA– CuP position	Position of veins of female tegmina	Number of stridulatory teeth	Shape of male cerci	Length of ovipositor /pronotum	Song structure	Syllables main part (20– 30°C) [ms]	Syllables main + additionalpart (20–30°C) [ms]	Melanism	Colouration of stripes at dorsolateral borders of pronotum	Distribution
I. rh. leonorae	1/2–2/3	±equal	±equal or slightly longer	obtuse	3/4–4/5	distinctly attached	intermediate	149–170	Fig. 94	>2.3	groups of syllables	50–>200 ms	– / 200–>400	absent	reddish	SW part of Rila–Rodopi Mountain group
I. rh. rhodopensis	1/2–2/3	±equal	±equal or slightly longer	obtuse	2/3–4/5	distinctly attached	intermediate	134–180	Fig. 95	>2.3	groups of syllables	100–>200	250–800	absent	reddish to brownish– black	W Rodopi Mts and neighbouring territories
I. rh. petkovi	1/2–3/4	±equal	±equal or slightly longer	obtuse	3/4–4/5	strongly attached	intermediate	140–>160	Fig. 96	2 –2.5	groups of 2–6 syllables	150–300	500–>2000	absent	brownish– black	E Rodopi Mts and neighbouring territories
I. modestior	1/2	wider	±equal or slightly longer	right	>3/4 –>4/5	very close to each other	reticulate	95–250	Fig. 97	>2.3	groups of syllables	150–350	280–>350	absent	reddish	C Bakans and C Europe
I. dobrogensis	1/2	wider	slightly shorter	right	3/4–4/5	very close to each other	reticulate	261–275	Fig. 98	2 –2.3	groups of elaborate syllables	300–420	730–1100	absent	reddish	Popina Island in Dobrogea ( Romania )
I. zubowskii	1/3–1/2	wider	±equal or slightly shorter	obtuse	3/4–4/5	very close to each other	reticulate	181–227	Fig. 99	>2.5	long sequence of syllables	180–250	290–430	absent	reddish	SW Ukraine , Moldavia , E and S Romania
I. brevicauda	1/3–1/2	wider	±equal or slightly longer	right	2/3–3/4	very close to each other	reticulate	200–220	See Heller et al. 2004: Fig. 36	<2	phrases of syllables	100–400	?	absent	reddish	Austria , Croatia , Slovenia
Carpathian
Basin (SK, PL,
long	UA, RO, HU)
I. aff. camptoxypha	1/3–1/2	narrower	shorter	obtuse	1/2–2/3	distinctly attached	reticulate	50–80	Fig. 100	±2	sequence of	25–42	<200	absent	reddish	reaching Austria ,
syllables	? Montenegro
and
? Macedonia
I. gulae	1/2	±equal	±equal or slightly longer	obtuse	2/3–3/4	distinctly attached	reticulate	100–130	Fig. 101	>2.3	isolated syllables	150–250	1500–2500	absent	reddish	Isolated in a forest of SE Bulgaria
Mountains of C
I. obtusa	1/2	±equal	±equal or slightly longer	obtuse	3/4–4/5	distinctly attached	reticulate	120–161	Fig. 102	>2.3	groups of syllables	100–450	250–>400	absent	reddish	and W Bulgaria and neighbouring
Serbia
reddish or
I. amplipennis	1/3–1/2	±equal or wider	1.3–1.5: 1	obtuse	2/3–3/4	widely separated	reticulate	72–112	Fig. 103	1.6–2	isolated elaborate syllables	>2000– 3400	–	present	masked with black in melanistic	The extreme SE Balkans, NW Anatolia
forms
I. speciosa	1/3–1/2	wider	slightly longer	obtuse	±3/4	widely separated	reticulate	154–170	Fig. 104	1.5–1.8	isolated elaborate syllables	>3500–>5000	–	present	reddish or masked with black in melanistic forms	S Romania , the Balkans (S to 40°), NW Anatolia

FIGURES 105–129. Female abdominal terminalia of the Balkan Isophya and I. yaraligozi (localities as in Figs 31–55 unless specifically given); figures of the whole ovipositor are followed by enlarged figure of the ovipositor base: 105— I. hospodar (BG: Dolni Glavanak, 25.04.2005, CC); 106— I. rectipennis (BG: Byala Vill., 27.06.2002, CC); 107— I. pavelii (BG: Chernogorovo, 29.05.2006, CC); 108— I. thracica ; 109— I. bureschi (BG: Bansko, 8.08.2006, CC); 110— I. yaraligozi ; 111— I. tosevski ; 112— I. andreevae (BG: Kresnensko Hanche, 19.04.2006, CC); 113— I. clara ; 114— I. miksici (BG: Gorski Dom Lodge, 24.06.2006, CC); 115— I. plevnensis (BG: Levishte, 14.07.2009, CC); 116— I. longicaudata adamovici (BG: Karandila, 24.06.2008, CC); 117— I. l. longicaudata (BG: Sofia University Botanical garden, 26.06.–1.07.2004, CC); 118— I. m. modesta (= intermedia ); 119— I. rhodopensis leonorae (BG: Livade, 9.08.2006, CC); 120— I. rh. rhodopensis (BG: Bachkovo, 600 m, 23.05.2004, CC); 121— I. rh. petkovi (BG: Gluhite Kamani, 23.06.2008, CC); 122— I. modestior (BG: Bankya, 24.05.2006, CC); 123— I. dobrogensis ; 124— I. zubowskii ; 125— I. aff. camptoxypha (data as in Fig. 75); 126— I. gulae ; 127— I. obtusa (BG: Ravnets Ridge, 14.07.2009, CC); 128— I. amplipennis (as in Fig. 78); 129— I. speciosa (BG: Bankya, 24.05.2006, CC). Indications on Fig. 127 show, respectively: G—gonangulum; L—lamella; P—lateral pit between lamella and gonangulum. Scale (if present) = 5 mm. A large number of taxa (i.e. presently 36 species) with various morphological features and song structures occur from Northwestern Anatolia to Central and Eastern Europe reaching the Altai Mountains in the East and the Pyrenees in the West. Warchałowska-Śliwa et al. (2008) classified the taxa studied by them in four groups: I. modesta , I. pyrenaea , I. costata and I. kraussii group. Yet, outlining these is complicated due to transitional morphological characters, undescribed songs, and great genetic similarity resulting in perplexed molecular phylogeny. The group is quite diverse morpho-acoustically but the transitions may be very smooth and thus clear distinction between the groups is frequently difficult. Excluding the majority of I. modesta group, making the transition between the primitive I. straubei and I. rectipennis groups and the typical Northern stock, most taxa concerned here have the following common morphological tendencies: (1) shortened tegmina both in males and females with blunt apical parts in females; (2) moderately or strongly approximated CuP and CuA; (3) female tegmina with reticulate venation; (4) lack of typical melanism or, in some taxa, tendency of development of melanistic colouration. Re-evaluating the known taxonomic characters we generally sustain the classification by Warchałowska-Śliwa et al. (2008), yet using the possibility of further subdivision of some of these groups. The following groups are considered here: (1) I. modesta group; (2) I. costata group; (3) I. kraussii group; (4) I. pyrenaea group.