A new species of tiger beetle of the Cicindela campestris group from southern Turkey, with remarks on the identity of C. herbacea Klug, 1832 and other taxa related to C. desertorum Dejean, 1825
Abstract
Franzen, Michael (2007): A new species of tiger beetle of the Cicindela campestris group from southern Turkey, with remarks on the identity of C. herbacea Klug, 1832 and other taxa related to C. desertorum Dejean, 1825. Spixiana 30 (1): 13-24, DOI: 10.5281/zenodo.16850914
Full text
13 SPIXIANA 30 1 13–24 München, 1. Mai 2007 ISSN 0341 – 8391 A new species of tiger beetle of the Cicindela campestris group from southern Turkey, with remarks on the identity of C. herbacea Klug, 1832 and other taxa related to C. desertorum Dejean, 1825 (Insecta, Coleoptera, Cicindelidae) Michael Franzen Franzen, M. (2007): A new species of tiger beetle of the Cicindela campestris group from southern Turkey, with remarks on the identity of C. herbacea Klug, 1832 and other taxa related to C. desertorum Dejean, 1825. – Spixiana 30/1: 13-24 Several populations of tiger beetles of the Cicindela campestris group from southern Turkey and Lebanon were investigated with respect to morphometric ratios, male genitalia, and color patterns. As a result, Cicindela herbacea from Lebanon appears to be most closely related to C. turkestanicoides and C. desertorum from eastern Turkey. The full species status of these taxa is provisionally kept but future research is needed to clear their relations. Populations of a taxon from southern Turkey previously referred to C. herbacea represent a new species, which is described as Cicindela thughurica, spec. nov. The new species appears to be more closely related to C. campestris, but differs from the latter by smaller relative aedeagus length, the presence of complete middle bands, and a brownish-green dorsal coloration in most specimens. Syntopic occurrence of C. herbacea and C. thughurica, spec. nov. is shown to occur in the northern Amanus Mountains, Turkey. Parapatric occurrence of the new species and C. campestris is known from the Bolkar daglari. Michael Franzen, Hauptstr. 1a, D-85467 Neuching, Germany Introduction The Cicindela campestris group of the Near East comprises the following taxa: C. campestris pontica Fischer, 1825, C. campestris palustris Motschulsky, 1840, C. campestris suffriani Loew, 1843, C. herbacea Klug, 1832, C. desertorum Dejean, 1825, C. turkestanicoides turkestanicoides Horn, 1938, C. turkestanicoides perreaui Deuve, 1987, and C. talyschensis Chaudoir, 1846 (e.g. Putchkov & Matalin 2003). The taxonomy of these taxa (belonging to “groupe V” sensu Rivalier 1950) in the Middle East is partly still insuffi ciently known. A major problem concerns the taxonomic status of the so-called Cicindela “herbacea” of southern Turkey, which specifi c status and relations are still under discussion (Korell 1988, 1994, Cassola 1999). Cicindela herbacea has been described from Lebanon (Klug 1832, see also Baker 1997). Subsequently, the name has been referred to various populations of green tiger beetles of the Near East (e.g. Israel, Turkey, “Armenia”, Cyprus: Horn & Roeschke 1891, Horn 1926, 1930, Mandl 1944, 1963, Nussbaum 1987, Korell 1988, Wiesner 1992, Korell 1994, Cassola 1999). Prior to Mandl (1944), who states only very shortly that the taxon requires a specifi c status, herbacea has been regarded as a subspecies of C. campestris. Most authors followed Mandl and treated herbacea as a separate species (e.g. Werner 1991, Wiesner 1992, Cassola 1999, Putchkov & Matalin 2003). However, its species status has been questioned by Korell (1988, 1994). The author mentioned several populations which appeared to be transitional between
14 C. campestris and C. herbacea and therefore proposed to regard herbacea only as a subspecies of C. campestris (Korell 1994). Finally, Cassola (1999) again recognized herbacea as a full species, because of signifi cant morphological differences to C. campestris (C. “herbacea” with a smaller, narrower head and pronotum and the presence of a complete, subtransversal, acutely bent middle band). However, Cassola also stated that campestris × herbacea intergrades may occur in the Antalya province of southwestern Turkey. During the last years I was able to collect series of all Anatolian taxa of the Cicindela campestris group, as well as to study material of Cicindela herbacea from Lebanon. The results of these investigations yield a different view on the taxonomy of the green tiger beetles of the genus Cicindela of the Levant and southern Turkey. Material and Methods Material of a Cicindela from southern Anatolia formerly referred to C. herbacea (e.g. Korell 1988, 1994, Cassola 1999) was compared to several morphologically similar taxa of the Cicindela campestris group of Turkey and the Levant. Since the taxonomy of the Near Eastern subspecies of C. campestris (nominal taxa C. campestris pontica, C. c. palustris, C. c. suffriani) urgently needs a revision, I use the term “C. campestris” for a morphotype occurring in the Taurus Mountains of Turkey, from the southern Aegean coast in the west to Erzincan province in eastern Anatolia (Franzen, unpublished data). “True C. herbacea” are represented by a sample from near Bcharré (northern Lebanon). Specimens from this locality perfectly match the illustration of the type specimen given by Klug (1832, pl. 21, fi g. 1), with respect to body proportions and elytral design. The female type of C. herbacea has been collected by C. G. Ehrenberg and W. F. Hemprich “in ora Syria prope Berytum”. According to Baker (1997: 171-172), Ehrenberg and Hemprich crossed the Lebanon Mountains twice, on the fi rst occasion near Djebel Sanin and on the second at Mount Lebanon/Bcharré. Since C. herbacea is supposed to be restricted to medium and high elevations (see below) the type is most probably originated from one of these localities. For statistical analysis, samples were aggregated as follows (see also Fig. 1 and Appendix for exact localities and collection data): “C. sp.” (Fig. 2): 71 specimens (39 males, 32 females) of so-called C. “herbacea” from scattered localities in southern and southeastern Anatolia; “Sertavul” (Fig. 3): 29 specimens (16 males, 13 females) from the area of the Sertavul pass in the central Taurus Mountains. Korell (1988, 1994) and Cassola (1999) mentioned C. “herbacea” or C. campestris × C. “herbacea” intergrades to occur at this locality; “Hasan” (Fig. 4): 25 specimens (18 males, 7 females) of C. campestris from the northeastern slope of the Hasan dag massif in Central Anatolia; “Bcharré” (Fig. 5): 17 specimens (10 males, 7 females) of C. herbacea from the area of Bcharré/Les Cedres at Mount Lebanon; “Zorkun” (Fig. 6): 18 specimens (10 males, 8 females) of C. cf. herbacea from the environments of Zorkun village in the northern Amanus mountains (Turkey); “Darbogaz” (Fig. 7): 20 specimens (10 males, 10 females) of C. cf. herbacea from the environments of Darbogaz village at the northern slope of the Bolkar daglari in the central Taurus Mountains; “Pontus” (Fig. 8): 40 specimens (19 males, 21 females) of C. desertorum from Ovit and Zigana passes in the eastern Pontus Mountains, northeastern Anatolia; “Van” (Fig. 9): 24 specimens (17 males, 7 females) of C. turkestanicoides perreaui (sensu Cassola 1999) from Karabet and Kuskun Kiran passes south of Lake Van, eastern Taurus Mountains, southeastern Anatolia. In addition, material of C. campestris from various localities in the Taurus Mountains west of the Sertavul pass (see Appendix for detailed locality data) was compared to the group “C. sp.”, with respect to their elytral markings and coloration. Measurements taken are: total length (TL, without labrum); head width (HW, across the widest point on the eyes); pronotum length (PL); maximum pronotum width (PWm); basal pronotum width (PWb); elytral length (EL, from scutellum to apex); humeral elytral width (EWh, width of left + right elytron at base, usually at the apical end of the humeral lunule); maximum elytral width (EWm, width of left + right elytron, usually at the middle band); aedeagus length (AL, in a straight line from apex to basal orifi ce); number of scapal setae (SS, on fi rst antenna segment, except apical sensories; mean of left and right antenna). In addition, the following ratios were calculated: HW/TL (relative head width); HW/PWm (relation of head width and maximum pronotum width); PL/TL (relative pronotum length); PL/PWm (relation of pronotum length and width); PWm/PWb (degree of lateral pronotum curvature); EL/TL (relative elytral length); EWm/TL (relative elytral width); EWm/ EL (relation of maximum elytral width and length); EWh/EWm (degree of lateral elytral curvature); AL/EWm (aedeagus length in relation to maximum elytral width). The above mentioned 10 morphometric ratios plus the characters TL and SS were analysed statistically. Because of possible sex dependent variation, analysis was run separately for males and females. Analysis was done using SPSS for Windows. Groups
15 mean values were z-transformed and Squared Euclidian distances were clustered using Ward’s method. Collection acronyms are: CFO (Coll. M. Franzen, Oberneuching), CHS (Coll. W. Heinz, Schwanfeld), ZSM (Zoologische Staatssammlung München). Results and Discussion 1. Morphological analysis Morphometry. According to the morphometric analysis (comparison of mean values), the investigated groups can be divided into two subgroups (Tabs 1, 2) which correspond to the two major clusters shown in Figure 10. One subgroup comprises the samples “C. sp.”, “Sertavul”, and “Hasan” (followingly named C. campestris subgroup = ca), the other comprises the samples “Bcharré”, “Zorkun”, “Darbogaz”, “Pontus”, and “Van” (C. desertorum subgroup = de). Distinguishing characters between these subgroups are that specimens of the campestris subgroup have broader pronota compared to head width (HW/PWm males ≤ 1.15 vs. ≥ 1.20 in de; females ≤ 1.13 vs. ≥ 1.15 in de), longer pronotal lengths in males (PL/TL 0.18 vs. ≤ 0.18 in de), posteriorly more narrowed pronota (PWm/PWb males ≥ 1.24 vs. ≤ 1.23 in de; females ≥ 1.28 vs. ≤ 1.29 in de), longer elytra in females (EL/TL ≤ 0.62 vs. ≥ 0.62 in de), broader bodies with less parallel-sided elytra (EWm/TL males ≥ 0.43 vs. ≤ 0.41 in de; females ≥ 0.43 vs. ≤ 0.43 in de; EWm/EL males ≥ 0.69 vs. ≤ 0.67 in de; females ≥ 0.70 vs. ≤ 0.69 in de), and shorter aedeagus lengths (AL/ EWm ≤ 0.79 vs. ≥ 0.90 in de) (see also Fig. 11). Within the C. campestris subgroup, “Sertavul” and “Hasan” samples are mostly identical or at least very weakly differentiated (characters and ratios TL, HW/TL, HW/PWm, PL/TL, EL/TL, EWm/TL, EWh/EWm, AL/EWm, SS in males; TL, HW/TL, HW/PWm, PL/TL, PL/PWm, EL/TL, EWh/EWm, SS in females). In contrast, the sample “C. sp.” seems to forms a sister group to the Sertavul/Hasan (= S/H) groups, with the “C. sp.” sample having smaller total lengths (TL males 10.9 vs. ≥ 12.4 in S/H; females 11.8 vs. ≥ 13.3 in S/H), broader heads (HW/TL males 0.27 vs. 0.26 in S/H; females 0.28 vs. 0.26 in S/H; HW/PWm males 1.15 vs. ≤ 1.10 in S/H; females 1.13 vs. ≤ 1.08 in S/H), less broader pronota (PL/PWm males 0.76 vs. ≤ 0.75 in S/H; females 0.71 vs. ≤ 0.70 in S/H), shorter elytra in males (EL/TL 0.61 vs. 0.62 in S/H), shorter aedeagus lengths (AL/EWm 0.79 vs. ≥ 0.81 in S/H; Fig. 11), and more scapal setae (SS males 2.4 vs. ≥ 3.4 in S/H; females 1.9 vs. ≥ 2.4 in S/H). Within the C. desertorum subgroup, differences between samples are weak and relations are partly sex depending. However, among the fi ve geographical samples, “Van” appears to be unique in possessing the highest mean values for males in the ratios HW/PWm (narrow pronota compared to head width), EL/TL (longest elytra), AL/EWm (longest aedeagi; Fig. 11), while females exhibit lowest means in EWm/TL and HW/TL (narrow elytral and head width). Both sexes of the “Van” sample possess lowest mean values in EWm/EL (narrow and elongate elytra). The remaining samples appear less differentiated since mean values largely overlap. In males, the southern samples “Bcharré”, “Zorkun”, “Darbogaz” are clustering cloth together and form a clade opposite to “Van”, based on similar or identical values in the ratios HW/PWm (relation of head width and maximum pronotum width), EL/TL, EWm/TL (elytral length and width in relation to total length), EWm/EL (elytral proportions), and AL/EWm (aedeagus length in relation to maximum elytral width). Moreover, there seems to be a clinal trend for the ratios HW/PWm (males + females), PL/PWm (females), and EWm/EL, with lowest values in the southernmost population (Bcharré), intermediates in Zorkun and highest values in the northernmost population (Darbogaz). Fig. 1. Collection sites of tiger beetles of the Cicindela campestris group used for statistical analysis. Large dots: 1 – Bcharré; 2 – Sertavul pass; 3 – Hasan dag; 4 – Darbogaz; 5 – Zorkun; 6 – Zigana pass (“Pontus”); 7 – Ovit pass (“Pontus”); 8 – Kuskun Kiran pass (“Van”); 9 – Karabet pass (“Van”). Unumbered small dots represent localities of “C. sp.” aggregated for analysis (see Fig. 13 for exact localities).
16 Tab. 2. Morphometrics of female tiger beetles of the C. campestris group. See “Material and Methods” for abbreviations. First line: mean ± standard deviation; second line (in parantheses): range. n TL HW/TL HW/PWm PL/TL PL/PWm PWm/PWb EL/TL EWm/TL EWm/EL EWh/EWm SS C. sp. 32 11.8±0.7 0.28±0 1.13±0.03 0.17±0 0.71±0.02 1.30±0.04 0.61±0.01 0.43±0.01 0.71±0.02 0.80±0.01 1.9±1.0 (10.8-13.2) (0.27-0.29) (1.06-1.19) (0.16-0.18) (0.66-0.74) (1.23-1.38) (0.59-0.63) (0.42-0.45) (0.68-0.75) (0.77-0.83) (0.5-4.0) Sertavul 13 13.3±0.6 0.26±0 1.08±0.03 0.17±0.01 0.70±0.03 1.28±0.03 0.62±0.01 0.43±0.01 0.70±0.02 0.79±0.01 2.4±1.2 (12.2-14.3) (0.25-0.26) (1.02-1.13) (0.16-0.17) (0.65-0.75) (1.24-1.34) (0.60-0.63) (0.42-0.45) (0.67-0.72) (0.77-0.81) 0-4.0) Hasan 7 13.6±1.0 0.26±0.01 1.04±0.02 0.17±0.01 0.70±0.03 1.32±0.07 0.61±0.01 0.45±0.01 0.73±0.03 0.78±0.02 3.4±2.2 (12.3-15.2) (0.25-0.27) (1.00-1.08) (0.17-0.18) (0.65-0.74) (1.23-1.44) (0.60-0.63) (0.42-0.46) (0.68-0.77) (0.75-0.81) (0-5.5) Bcharré 7 12.3±0.6 0.27±0 1.15±0.03 0.16±0 0.67±0.02 1.27±0.02 0.63±0.01 0.43±0.01 0.69±0.01 0.79±0.02 3.2±1.7 (11.4-13.4) (0.26-0.27) (1.09-1.19) (0.15-0.16) (0.65-0.70) (1.24-1.30) (0.62-0.64) (0.43-0.44) (0.67-0.70) (0.77-0.81) (0-5.0) Zorkun 8 12.9±0.7 0.26±0.01 1.16±0.02 0.17±0.01 0.70±0.01 1.29±0.03 0.63±0.01 0.42±0.01 0.67±0.02 0.78±0.01 2.4±0.4 (11.8-14.2) (0.25-0.27) (1.13-1.18) (0.16-0.18) (0.69-0.72) (1.25-1.36) (0.61-0.64) (0.40-0.44) (0.64-0.70) (0.76-0.79) (2.0-3.0) Darbogaz 10 13.1±0.6 0.26±0 1.20±0.02 0.17±0.01 0.71±0.03 1.28±0.02 0.62±0.01 0.42±0.01 0.67±0.02 0.79±0.02 3.6±1.7 (12.2-14.2) (0.25-0.26) (1.15-1.23) (0.16-0.18) (0.68-0.78) (1.26-1.32) (0.61-0.64) (0.40-0.45) (0.64-0.71) (0.74-0.84) (1.5-6.5) Pontus 21 14.1±0.6 0.26±0.01 1.21±0.04 0.18±0.01 0.72±0.03 1.28±0.05 0.62±0.02 0.42±0.02 0.68±0.02 0.80±0.02 2.7±1.4 (12.6-15.1) (0.24-0.29) (1.16-1.30) (0.16-0.20) (0.66-0.77) (1.17-1.38) (0.60-0.71) (0.40-0.47) (0.65-0.71) (0.78-0.84) (0.5-5.0) Van 7 14.5±0.4 0.25±0.01 1.18±0.05 0.16±0 0.72±0.02 1.27±0.07 0.63±0.01 0.41±0.01 0.64±0.01 0.79±0.01 4.2±2.5 (13.8-14.9) (0.24-0.26) (1.10-1.26) (0.16-0.17) (0.69-0.75) (1.20-1.40) (0.61-0.65) (0.39-0.41) (0.62-0.65) (0.78-0.82) (2.0-9.0) Tab. 1. Morphometrics of male tiger beetles of the C. campestris group. See “Material and Methods” for abbreviations. First line: mean ± standard deviation; second line (in parantheses): range. n TL HW/TL HW/PWm PL/TL PL/PWm PWm/PWb EL/TL EWm/TL EWm/EL EWh/EWm AL/EWm SS C. sp. 39 10.9±0.6 0.27±0.01) 1.15±0.04 0.18±0.01 0.76±0.03 1.25±0.04 0.61±0.01 0.43±0.01 0.70±0.02 0.80±0.01 0.79±0.03 2.4±1.3 (9.8-11.8) (0.26-0.29 (1.09-1.24) (0.17-0.19) (0.70-0.81) (1.17-1.34) (0.58-0.62) (0.41-0.44) (0.67-0.74) (0.76-0.83) (0.72-0.86) (0-5.0) Sertavul 16 12.7±0.4 0.26±0.00 1.10±0.03 0.18±0.01 0.75±0.02 1.24±0.05 0.62±0.01 0.43±0.01 0.69±0.03 0.80±0.03 0.81±0.03 3.4±2.4 (11.8-13.5) (0.25-0.27) (1.06-1.15) (0.17-0.19) (0.73-0.81) (1.14-1.33) (0.60-0.65) (0.41-0.47) (0.64-0.76) (0.74-0.83) (0.76-0.86) (0-8.0) Hasan 18 12.4±0.4 0.26±0.01 1.06±0.03 0.18±0 0.72±0.03 1.26±0.04 0.62±0.01 0.44±0.01 0.71±0.02 0.80±0.02 0.83±0.03 5.1±2.5 (11.7-12.9) (0.25-0.28) (1.00-1.13) (0.17-0.19) (0.67-0.77) (1.19-1.35) (0.60-0.63) (0.42-0.46) (0.66-0.75) (0.74-0.84) (0.80-0.89) (0.5-11.0) Bcharré 10 12.0±0.4 0.26±0.01 1.20±0.04 0.16±0.01 0.72±0.03 1.23±0.03 0.62±0.01 0.41±0.01 0.67±0.01 0.80±0.01 0.92±0.03 2.5±1.0 (11.5-12.8) (0.25-0.26) (1.16-1.25) (0.16-0.17) (0.68-0.76) (1.18-1.29) (0.60-0.64) (0.40-0.43) (0.65-0.69) (0.78-0.82) (0.88-0.98) (1.0-4.0) Zorkun 10 11.8±0.2 0.26±0.01 1.21±0.03 0.17±0.01 0.77±0.04 1.19±0.03 0.62±0.01 0.41±0.01 0.66±0.02 0.79±0.02 0.98±0.03 2.6±1.1 (11.5-12.3) (0.25-0.26) (1.16-1.29) (0.16-0.18) (0.71-0.83) (1.11-1.23) (0.60-0.63) (0.40-0.42) (0.64-0.69) (0.77-0.84) (0.93-1.03) (1.0-5.0) Darbogaz 10 12.1±0.4 0.25±0.01 1.23±0.03 0.17±0 0.75±0.03 1.20±0.03 0.62±0.01 0.41±0.01 0.65±0.02 0.79±0.02 1.0±0.03 4.1±1.5 (11.4-12.6) (0.24-0.26) (1.19-1.27) (0.16-0.18) (0.70-0.79) (1.15-1.24) (0.61-0.64) (0.40-0.42) (0.63-0.68) (0.77-0.84) (0.94-1.03) (1.5-6.0) Pontus 19 13.3±0.5 0.26±0.01 1.24±0.04 0.18±0 0.76±0.03 1.23±0.06 0.61±0.01 0.41±0.01 0.67±0.02 0.82±0.02 0.90±0.03 3.7±1.5 (12.0-13.8) (0.24-0.27) (1.14-1.31) (0.17-0.19) (0.69-0.81) (1.12-1.36) (0.60-0.62) (0.39-0.43) (0.64-0.72) (0.79-0.85) (0.87-0.99) (1.0-6.3) Van 17 13.4±0.7 0.25±0.01 1.25±0.03 0.17±0.01 0.77±0.03 1.20±0.04 0.63±0.01 0.40±0.01 0.64±0.02 0.79±0.02 1.04±0.04 3.4±1.8 (12.5-14.6) (0.24-0.26) (1.19-1.32) (0.16-0.18) (0.69-0.81) (1.14-1.26) (0.61-0.64) (0.38-0.42) (0.61-0.67) (0.76-0.83) (0.97-1.10) (0-7.0)
17 Figs. 2-9. Individuals of the Cicindela campestris group from Turkey and Lebanon representing the samples of the morphometric analysis (females). 2. C. thughurica, spec. nov. (“C. sp.”; near Kaypak, 680 m, Osmaniye prov., Turkey). 3. C. campestris (“Sertavul”; Sertavul pass, 1500 m, Içel prov., Turkey). 4. C. campestris (“Hasan”; Hasan dag, 1500-1600 m, Aksaray prov., Turkey). 5. C. herbacea (“Bcharré”; Bcharré/Les Cedres, Lebanon). 6. C. herbacea (“Zorkun”; near Zorkun village, 1750-1850 m, Osmaniye prov., Turkey). 7. C. herbacea (“Darbogaz”; above Darbogaz village, 1600-1900 m, Nigde prov., Turkey). 8. C. desertorum (“Pontus”; Ovit pass, 2600 m, Rize prov., Turkey). 9. C. turkestanicoides perreaui (“Van”; Kuskun Kiran pass, 2200-2300 m, Van prov., Turkey). 2345 6789 Coloration. Under diffuse natural light, the dorsal coloration of all groups, with the exception of “C. sp.”, is metallic green, with more or less distinctive coppery to reddish patches and refl ections on the head, pronotum and along the sutural and lateral margins of the elytra. In contrast, individuals of the “C. sp.” group mostly exhibit a brownish green or dull olive dorsal coloration under diffuse natural light. However, colors of “C. sp.” also appear clearly metallic greenish under strong direct light.
18 Elytral markings: Specimens of all samples possess an interrupted humeral lunule, and mostly a complete apical lunule (Figs. 2-9). A complete middle band is present in all specimens of the groups “Bcharré”, “Zorkun”, “Darbogaz”, “Pontus”, and “Van” (Figs. 5-9). It is mostly not interrupted in the “C. sp.” sample (Fig. 2), with the exception of fi ve individuals (= 6.6 %) which have the middle band interrupted at least on one elytron. In contrast, the groups “Sertavul” and “Hasan” exhibit mostly interrupted middle bands (Figs. 3, 4) with eight individuals from “Sertavul” and seven from “Hasan” (= 28 % each) having the middle bands not interrupted. Three individuals of comparative C. campestris from Aktoprak and Karagöl, localities which are geographically nearest to the “C. sp.” sample (approximately 15-30 km: Figs. 1, 13), have all interrupted middle bands (Figs. 14). Among the comparative material of C. campestris from the western Taurus Mountains (see Appendix) individuals from two localities also exhibit complete middle bands. Those individuals make 50 % of the Davraz dag sample (6 of 12) and 20 % of a small series from Irmasan pass (1 of 5). In contrast, all specimens from Usak province, Yatagan, Gökçeören, Keçiborlu, Phaselis, and Alanya (n = 49) possess all broadly interrupted middle bands. The “Van” sample is unique in possessing narrow and strongly oblique middle bands (Fig. 9). Shape of aedeagus: The outer shape of the male genitalia is rather similar within the two subgroups. Specimens of the desertorum subgroup (“Bcharré”, “Darbogaz”, “Zorkun”, “Van”, “Pontus”: Figs. 12 C-F) exhibit a very long and elongate aedeagus, while it is comparably stout in members of the campestris subgroup (“C. sp.”, “Sertavul”, “Hasan”: Figs. 12 A-B). 2. Conclusions Summarizing the results of the morphological analyses the following conclusions can be drawn: 1. Based on the morphological data, the Middle Eastern species of the “groupe V” sensu Rivalier (1950) can be provisionally arranged in two subgroups, namingly the Cicindela campestris subgroup (containing C. campestris and a new species described below) and the Cicindela desertorum subgroup (containing C. desertorum, C. turkestanicoides, and C. herbacea; C. talyschensis also tentatively included). In the Middle East, members of the C. desertorum subgroup are usually restricted to humid, medium and high elevations (1000-3000 m a.s.l.), especially within dry continental or Mediterranean climates. In contrast, members of the C. campestris subgroup inhabit large altitudinal ranges from sea level to high elevations. 2. Two samples from southern Turkey (“Darbogaz”, “Zorkun”) are morphologically almost identical with C. herbacea from Lebanon and are referable to this taxon. Slight differences between the samples may indicate an infraspecifi c variation. Fig. 10. Phenetic relationships between the investigated groups based on morphometric ratios (mean values; squared Euclidian distances, clustered using Wards’s method). Upper: males; lower: females.
19 3. Cicindela herbacea appears to be most similar to C. desertorum and especially C. turkestanicoides. It should be noted that Klug (1832) already mentioned the similarity of his new species and C. desertorum in the fi rst sentence of his diagnosis of C. herbacea (“C. desertorum Boeb. proxima ...”). However, at present it seems to be appropiate to treat C. herbacea and C. turkestanicoides provisionally as separate species, although distinguishing characters are weak. Future researches should clear the relations of all taxa involved, based on more extensive geographic samples (including topotypical material of C. t. turkestanicoides from northeastern Iran; type locality: “Taesch und Schaku (zwischen Astrabad und Scharud)” [= between Gorgan and Emâmrûd, eastern Elburz Mountains]) and the analysis of more characters (e.g., female genitalia, males’ inner sac structures, molecular data sets). 4. The groups “Hasan” and “Sertavul” are identical in most morphological characters and both must be referred to C. campestris. 5. Specimens from southern Turkey formerly referred to C. “herbacea” (= C. sp.) are morphologically different from “true” C. herbacea from Lebanon. Specimens of this form appear to be most similar to C. campestris but can not be attributed to any of the investigated forms, and therefore represent an undescribed taxon. The specifi c status of this form is supported by syntopic occurrence with C. herbacea and close parapatric occurrence with C. campestris in southern Turkey (see below). Cicindela thughurica, spec. nov. Figs 2, 12B, 15 Cicindela herbacea, Mandl 1963: 45 (localities “Namrun”, “Pozanti”). Cicindela herbacea, Korell 1988: 100 (part.: localities “Osmaniye: Karatepe”, “Amanos Dagl.: Akbes”, “Pozanti und Namrun”). Cicindela herbacea (sp. inqu.), Korell 1994: 43 (part.: localities “Pozanti [Adana]”, “nördlich Maras [Kahramanmaras]”). Cicindela herbacea, Cassola 1999: 238 (part.: localities “Namrun”, “Adana”, “Pozanti”, “Osmaniye, Karatepe”, “Osmaniye”, “N of Maras”, “Akbez”, “Gülek”, “Alexandrette”). Types. Holotype: M, with the following labels: “TR, Prov. Osmaniye: Strassenkr. Ri. Kaypak (nw. Fevsipasa), 680 m. Feuchte, lehmige Böschung. 08.04.1998, Franzen & Gruber leg.” [typed and printed white label with black margin], “3” [handscript round label], “Cicindela thughurica Franzen, 2007, Holotypus” [typed and printed red label] (in ZSM). – Paratypes. A total of 76 specimens, all from Turkey. 14MM and 8WW with the same data as the holotype (1M, 1W ZSM, 13MM, 7WW in CFO); 1M, 1W: above Zorkun (Osmaniye prov.), 17501850 m, 18.6.1997, Franzen leg. (CFO); 2MM, 1W: above (E of) Yarpuz (Osmaniye prov.), 1550 m, 19.6.1997, Franzen leg. (CFO); 3MM, 4WW: 5 km N Hieropolis-Castabala (Osmaniye prov.), 150 m, 9.4.1997, Franzen leg. (CFO); 4MM, 2WW: environments of Çiftehan (Nigde prov.), 850 m, 3.4.1988, de Freina leg. (5 CHS, 1 CFO); 1M, 2WW: eastern slope of Karahan pass (Malatya prov.), 1300 m, 30.4.1999, Franzen leg. (CFO); 5MM, 2WW: N of Tekir (Kahramanmaras prov.), 1100 m, 7.4.1998, Franzen & Gruber leg. (CFO); 9MM, 7WW: 30 km NW Kahramanmaras (Kahramanmaras prov.), 580 m, 7.4.1998, Franzen & Gruber leg. (CFO); 1W: environments of Kahramanmaras (“Marasch, Syrien”, Kahramanmaras prov.), Reitter (CFO); 1M: between Çiftehan and Pozanti (Adana prov.), 900-1000 m, 17.4.1973, Heinz leg. (CFO); 1M, 1W: E of Pozanti (Adana prov.), 800 m, 11.4.1998, Franzen leg. (CFO); 1W: Pozanti (Adana prov.), 25.5. 1961, Cadamuro leg. (CHS); 1W: Çatalan (“Anatolia, Prov. Adana, Catalon”, Adana prov.), 50100 m, 18.4.1985, Barries leg. (CFO); 1M, 2WW: E of Çamliyayla (Içel prov.), 1100 m, 11.4.1998, Franzen leg. (CFO); 1W: environments of Akbez (“Akbez, Syr.”, Gaziantep prov.), Winkler (CFO). Type locality. Turkey, Osmaniye Province: at Kaypak road near to the junction with the old Osmaniye-Fevsipasa road at 37°09.76'N, 36°28.00'E, 680 m elevation. Referred specimen: 1W: “Libaah, Syrien”, without further collection data (CFO). Diagnosis. A member of the Cicindela campestris group (= “groupe V” sensu Rivalier 1950), with a unique character combination among Middle Eastern species: small relative aedeagus length (≤ 3.8 mm, AL/EWm ≤ 0.86), small total length (≤ 11.8 mm in males, ≤ 13.2 mm in females), brownish-green dorsal 3.0 3.5 4.0 4.5 5.0 5.5 6.0 4.0 4.5 5.0 5.5 6.0 Maximum elytral width (mm) Aedeagus length (mm) C. sp. Sertavul Hasan Bcharré Zorkun Darbogaz Pontus Van Fig. 11. Relation of aedeagus length and elytral width within the investigated groups.
20 coloration and complete middle bands in most specimens. C. thughurica, spec. nov. differs from Near Eastern C. campestris by small aedegus length (≤ 3.9 mm vs. ≥ 4.1 mm), by less reduced elytral markings (middle bands complete in most C. thughurica, interrupted in most C. campestris), and by elytral coloration (brownish-green in C. thughurica, green in C. campestris). C. thughurica, spec. nov. differs from similar and geographic nearest (partly sympatric) taxa C. herbacea, C. desertorum, and C. turkestanicoides by its smaller relative aedeagus length (AL/EWm 0.72-0.86 vs. 0.87-1.10), by the more robust and less elongate shape of the aedeagus with a less pronounced apex, by its smaller total lengths, and a different elytral coloration (mostly brownish-green in C. thughurica, green in C. desertorum and C. turkestanicoides). In addition, C. thughurica, spec. nov. differs from C. talyschensis of northern Iran and Azerbaidshan in possessing a setose frons (glabrous in C. talyschensis) and in males by the rounded lateral sides of the elytra (elytral margins almost parallel in male C. talyschensis). Description of the Holotype M, total length (without labrum) 10.6 mm. Head width 2.8 mm (27 % of TL); head distinctly broader than pronotum (HW/PWm 1.12). Second antennal segments with 3 terminal (sensory) and 4 scapal setae each, third segment glabrous, fourth with 9 erected setae each, fi fth with 3 erected setae each. Anntenal segments 6-12 with very short, fi ne hairs. Mandibles with four teeth. Labrum broad, with 6 submarginal setae (right outermost broken), without medial tooth. Clypeus and genae glabrous. Frons and vertex of head with abundant, long, erect setae in addition to some pairs of supraorbital setae. Vertex strongly concave between eyes, surface strongly rugose, rugae forming fi ne, parallel ridges near eyes. Pronotum transverse (length 1.9 mm [= 18 % of TL], maximum width 2.5 mm; PL/PWm 0.76), anteriorly distinctly wider than posteriorly (basal pronotum width 2.1 mm; PWm/PWb 1.20). Pronotum deeply wrinkled at lateral margins, shallowly wrinkled on disc, with numerous long setae; anterior transverse sulcus deep, V-shaped; posterior transverse sulcus deep, W-shaped; median longitudinal sulcus shallow. Proepisterna, hindepisterna, and lateral parts of the metasternum with long white setae. Median metasternum glabrous. Abdominal sternites 3-6 with numerous scattered short, fi ne setae. Procoxa with a small group of long, erect setae, mesocoxa with a large central spot of dense, long, erect setae, and metacoxa with a few scattered fi ne erect setae. Elytral length 6.5 mm (= 61 % of TL); humeral elytral (body) width 3.6 mm; maximum elytral (body) width at middle band (EWm) 4.5 mm (= 42 % of TL, 69 % of EL). Elytral surface densely scattered with numerous small pits, each associated with a small smooth granule. Granules are most conspicuous at base and less distinctly apically. Elytral apex without microserrulations; small apical spine present. Front and middle trochanters with one subapical seta on each. Front and middle femora dorsally and ventrally with numerous long white erect setae, hind femora only ventrally with setae. Tibia with short white, erect setae. Tarsi with very short white setae. Aedeagus (Fig. 12 B): length 3.6 mm (ratio AL/EWm 0.81). Coloration: Anntenal segments 1-5 metallic red, Fig. 12. Aedeagi of green tiger beetles of Turkey and Lebanon (scale bar represents 1 mm): A, Cicindela campestris (Sertavul pass, Içel prov., Turkey); B, C. thughurica, spec. nov., holotype (“C. sp.”, near Kaypak, Osmaniye prov., Turkey); C, C. herbacea (Bcharré, Lebanon); D, C. herbacea (above Darbogaz, Nigde prov., Turkey); E, C. turkestanicoides perreaui (Kuskun Kiran pass, Van prov., Turkey); F, C. desertorum (Ovit pass, Rize prov., Turkey). A B C D E F
21 remainder blackish. Mandibles bicolored, with teeth black and remainder brownish-yellow. Labrum brownish-yellow, with a fi ne black anterior margin. Clypeus and genae both metallic red with greenish margins. Head and pronotum dorsally dull coppery to red with greenish refl ections (overall appearence brownish under diffuse daylight). Elytra metallic greenish with coppery to red refl ections, elytral pits bluish (overall elytral coloration appears brownish under diffuse daylight). A diffuse, dark brownish area around the inner (sutural) part of the middle band. Elytral lateral margins and suture coppery to red. Elytral maculations creamish-white. Humeral lunule divided, broken into a humeral and posthumeral dot. Marginal band lacking, middle band complete, not broken into marginal and inner dots. Apical lunule complete, although bands between apical and subapical parts are very thin. Anterior underside (proepisterna, hindepisterna, and lateral parts of metasternum) metallic reddish, posteriorly (coxa, median metasternum, abdominal sternites 3-6) dark metallic violet to almost black. Trochanters black, femora metallic coppery to reddish, tibia Fig. 13. Distribution of tiger beetles of the Cicindela campestris group in central southern Turkey (based on material proofed by the author). %: C. thughurica, spec. nov.; #: C. herbacea; Œ: syntopic C. thughurica and C. herbacea; ’: C. campestris; ! : C. turkestanicoides perreaui; ª : syntopic/sympatric C. thughurica, spec. nov. and C. turkestanicoides perreaui. 1 – Sertavul pass; 2 – Aktoprak; 3 – Karagöl; 4 – Hasan dag; 5 – Göreme ; 6 – Erciyes dagi; 7 – Darbogaz; 8 – Zorkun; 9 – Çamliyayla; 10 – Pozanti; 11 – Ciftehan; 12 – Çatalan; 13 – Hieropolis-Castabala; 14 – Kaypak; 15 – Yarpuz; 16 – Akbez; 17 – Kahramanmaras; 18 – 30 km NW Kahramanmaras; 19 – Tekir; 20 – Karahan pass; 21 – Kubbe pass. metallic greenish, tarsi dull metallic greenish with violet refl ections. Variation. Variation of morphometric characters is shown in Tabs 1 and 2 (“C. sp.”). In contrast to the holotype, a small, black labral medial tooth is present in most specimens of the paratypes. Distribution. To date, C. thughurica is only known from southern Turkey (Fig. 13). Localties range from the southern slopes of the Bolkar daglari (Çamliyala, Pozanti) in the west to the Karahan pass (west of Malatya) in the northeast. Southernmost reliable localities are in the central Amanus Mountains (Osmaniye prov.). However, Cassola (1999) mentioned C. “herbacea” from Iskenderun, Hatay prov. (“Alexandrette, Asia minor”, Kricheldorff), a locality at the western base of the southern Amanus mountains. Most probably, this record also refers to C. thughurica. Korell (1988, 1994) and Cassola (1999) list some localities of C. “herbacea” from the western Taurus Mountains, west of Sertavul pass (e.g., Irmasan pass, Akseki, Egridir, Gazipasa, Aphrodisias).