A review of Eupholidoptera (Orthoptera, Tettigoniidae) from Crete, Gavdos, Gavdopoula, and Andikithira Author Willemse, Luc https://orcid.org/0000-0003-0517-9778 Naturalis, PO Box 9517, 2300, RA Leiden, Netherlands luc.willemse@naturalis.nl Author Tilmans, Jos https://orcid.org/0000-0003-0127-6191 Herkenboscher Strasse 33, 41849, Wassenberg-Rothenbach, Germany Author Kotitsa, Nefeli Natural History Museum Crete, University of Crete, P. O. Box 2208, 71409, Heraklion, Crete, Greece & Institute of Biodiversity and Ecosystem Research, Bulgarian Academy of Sciences, 1 Tsar Osvoboditel blvd., 1000, Sofia, Bulgaria Author Trichas, Apostolos https://orcid.org/0000-0002-7917-5262 Natural History Museum Crete, University of Crete, P. O. Box 2208, 71409, Heraklion, Crete, Greece Author Heller, Klaus-Gerhard https://orcid.org/0000-0002-3331-3228 Triesdorf Bahnhof 8, 91732, Merkendorf, Germany Author Chobanov, Dragan https://orcid.org/0000-0002-1642-0363 Institute of Biodiversity and Ecosystem Research, Bulgarian Academy of Sciences, 1 Tsar Osvoboditel blvd., 1000, Sofia, Bulgaria Author Ode, Baudewijn https://orcid.org/0000-0002-8929-2737 Riethorsterweg 12, 6586, AC Plasmolen, Netherlands text ZooKeys 2023 2023-03-01 1151 67 158 http://dx.doi.org/10.3897/zookeys.1151.97514 journal article http://dx.doi.org/10.3897/zookeys.1151.97514 1313-2970-1151-67 5FEDE55DC9AF47D591259F1758AE2A18 0B3CEF4A954955478878AD15C022DCD5 Eupholidoptera annamariae Nadig, 1985 Figs 18 , 32 , 46 , 60 , 76 , 90 , 104 , 118 , 119 , 133 , 147 , 161 , 175 , 190 , 206 , 240 , 254 , 256 , 259 Eupholidoptera annamariae Nadig 1985 : 329. Eupholidoptera annamariae Morphological description. Nadig 1985 : 329. Eupholidoptera annamariae Bioacoustics. Ciplak et al. 2009 : 27, 54. Examined specimens. 1 ♂, 1 ♀ ( paratypes ); 23 ♂, 24 ♀ (for details see Suppl. material 2). Diagnostic features. Frontal part of head (Fig. 18 ) pale with black dots; frontal half of pronotal disc (Fig. 32 ) predominantly black sharply transversely delineated from pale rear half. Male (Fig. 240 ) - stridulatory file with 109 teeth (89 in Nadig 1985 ) (including proximal and distal ones), density of teeth in middle two thirds of the file 19 teeth per mm; anal tergite (Figs 76 , 90 , 104 ) with hind margin medially strongly bent downward forming two very long, curved teeth covered with small denticles pointing downward and slightly outward; cerci (Figs 118 , 133 ) unarmed, 5 x longer than wide, basal half cylindrical, apical half conical, slightly curved inward halfway subtly widened, in profile straight; subgenital plate (Figs 147 , 161 ) wider than long, widest halfway, sides widely rimmed, in profile strongly upturned, pointing upward, tip apical lobes rounded, spineless, excised along one third of length; styli (Fig. 175 ) short, ca. one third as long as cerci, 1.5-2.0 x longer than wide, inserted ventrally just proximal of tip of apical lobe, pointing downward; titillator (Figs 190 , 206 ) slightly asymmetrical, apical arms strong, narrow, fused along entire length, smooth, evenly curved, apically narrowed toward needle shaped tip, pointing somewhat left or right. Female - subgenital plate (Figs 46 , 60 ) circular to transverse oval with central longitudinal hump, adjoined by deep grooves, apical lobes touching, hind margin medially excised along one quarter to one third of length, in profile deltoid, with deep ventral groove, tip obtuse angular. Figures 11-24. Colour pattern of head, frontal view 11 Eupholidoptera smyrnensis ♀ Makrigiannis RMNH5014918 12 Eupholidoptera gemellata ♂ Mt. Idhi RMNH.INS1141843 13 Eupholidoptera pallipes ♂ paratype Mt. Lefka RMNH.INS1105313 14 Eupholidoptera cretica ♂ Mt. Lefka, Omalos RMNH.INS114838 15 Eupholidoptera latens ♀ Mt. Lefka, Kalergi CT1987.047.04 16 Eupholidoptera giuliae ♂ Argoules CT1995.011.18 17 Eupholidoptera francisae sp. nov. ♀ allotype Andikithira CT 2002.004.11 18 Eupholidoptera annamariae ♀ Kato Zakros CT1995.020.03 19 Eupholidoptera astyla ♀ Ano Viannos CT1987.024.03 20 Eupholidoptera feri ♂ holotype Mt. Dikti, Katharo plain RMNH.INS1105297 21 Eupholidoptera mariannae ♀ Ag. Ioannis RMNH.5014906 22 Eupholidoptera forcipata ♂ Mt. Idhi CT1987.044.03 23 Eupholidoptera marietheresae sp. nov. ♂ paratype Mt. Dikti, FC1606 CT2000.096.01 24 Eupholidoptera jacquelinae ♂ holotype Gavdos CT2000.005.02. Scale bars: 5 mm. Figures 25-38. Colour pattern of pronotum, dorsal view 25 Eupholidoptera smyrnensis ♂ Makrigiannis RMNH.INS152909 26 Eupholidoptera gemellata ♂ Mt. Idhi, Amariou RMNH.INS1141843 27 Eupholidoptera pallipes ♂ paratype Mt. Lefka RMNH.INS1105313 28 Eupholidoptera cretica ♂ Mt. Lefka, Omalos RMNH.INS1141838 29 Eupholidoptera latens ♂ Rhodopos CH8236 30 Eupholidoptera giuliae Chora Sfakion ♂ CT2000.014.10 31 Eupholidoptera francisae sp. nov. ♂ holotype Andikithira CT2002.004.04 32 Eupholidoptera annamariae ♂ Kato Zakros CT2000.030.04 33 Eupholidoptera astyla ♂ Krotos RMNH.INS1141819 34 Eupholidoptera feri ♂ holotype Mt. Dikti, Katharo plain RMNH.INS1105297 35 Eupholidoptera mariannae ♀ Kalavros RMNH.5086974 36 Eupholidoptera forcipata ♂ Mt. Idhi CT1987.044.03 37 Eupholidoptera marietheresae sp. nov. ♂ holotype Mt. Dikti RMNH.INS1141850 38 Eupholidoptera jacquelinae ♂ holotype Gavdos CT2000.005.02. Scale bars: 5 mm. Figures 39-52. Female subgenital plate in ventral view 39 Eupholidoptera smyrnensis Makrigiannis RMNH.5014918 40 Eupholidoptera gemellata Mt. Idhi, FC1602 CT2000.095.02 41 Eupholidoptera pallipes allotype Mt. Lefka RMNH.INS1105312 42 Eupholidoptera cretica Mt. Lefka, Omalos RMNH.INS1141837 43 Eupholidoptera latens Mt. Lefka, Kalergi CT1987.047.04 44 Eupholidoptera giuliae Argoules CT1995.011.03 45 Eupholidoptera francisae sp. nov. allotype Andikithira CT2002.004.11 46 Eupholidoptera annamariae Kato Zakros CT1995.020.03 47 Eupholidoptera astyla Krotos RMNH.INS1141820 48 Eupholidoptera feri allotype Mt. Dikti, Katharo plain RMNH.INS1105298 49 Eupholidoptera mariannae Kalavros RMNH5014912 50 Eupholidoptera forcipata Mt. Idhi, FC1602 RMNH.INS1141845 51 Eupholidoptera marietheresae sp. nov. allotype Mt. Dikti RMNH.INS1141849 52 Eupholidoptera jacquelinae Gavdos allotype CT2001.004.13. Scale bars: 1 mm. Figures 53-66. Female subgenital plate in lateral view 53 Eupholidoptera smyrnensis Makrigiannis RMNH.5014918 54 Eupholidoptera gemellata Mt. Idhi, FC1602 CT2000.095.02 55 Eupholidoptera pallipes allotype Mt. Lefka RMNH.INS1105312 56 Eupholidoptera cretica Mt. Lefka, Omalos RMNH.INS1141837 57 Eupholidoptera latens Mt. Lefka, Kalergi CT1987.047.04 58 Eupholidoptera giuliae Argoules CT1995.011.03 59 Eupholidoptera francisae sp. nov. allotype Andikithira CT2002.004.11 60 Eupholidoptera annamariae Kato Zakros CT1995.020.03 61 Eupholidoptera astyla Krotos RMNH.INS1141820 62 Eupholidoptera feri allotype Mt. Dikti, Katharo plain RMNH.INS1105298 63 Eupholidoptera mariannae Kalavros RMNH5014912 64 Eupholidoptera forcipata Mt. Idhi, FC1602 RMNH.INS1141845 65 Eupholidoptera marietheresae sp. nov. allotype Mt. Dikti RMNH.INS1141849 66 Eupholidoptera jacquelinae allotype Gavdos CT2001.004.13. Scale bars: 1 mm. Figures 67, 68. Female subgenital plate in lateral view 67 Eupholidoptera francisae sp. nov. Anidhroi CT2001.002.07 68 Eupholidoptera latens Prases DC-Ort000512. Measurements. See Tables 6 , 7 . Bioacoustics. Based upon the sound recordings of 6 specimens (53 syllables measured), the song of E. annamariae , as in all species of Eupholidoptera , consists of isolated syllables produced in long series with the opening hemisyllable much shorter and weaker than the closing hemisyllable. In E. annamariae , the syllable duration is ~ 160 ms. In the present recordings, the syllable repetition rate is very low. Published records ( Ciplak et al. 2009 ) show a syllable duration of ~ 128 ms at 25 °C and a syllable repetition rate of 1/s at maximum. The song may most likely be confused with the other species of Eupholidoptera in Crete, except E. smyrnensis and E. forcipata . For details of sound recordings of Eupholidoptera annamariae see Suppl. material 3. Variation. The colour pattern and genitalia in E. annamariae in the specimens studied show little variation with one exception. In one of four males collected along the northern coast of Lasithi, west of Sitia, near Xerokampos the cerci showed a clearly developed inner tooth, halfway the cercus (Fig. 119 ). Other morphological traits in this aberrant male as well as in the other three males from Xerokampos fitted E. annamariae . The cercus with a side tooth is considered an anomaly. It is noteworthy that the location where the aberrant male of E. annamariae was found is the northwestern most location of E. annamariae only some 5 km away from Kalavros where E. mariannae was found. Differential diagnosis. Males differ from congenerics in the anal tergite (Figs 76 , 90 , 104 ) with the uniquely shaped long, curved spined teeth pointing downward and inward, in the stout, unarmed almost straight cercus (Figs 118 , 133 ), in the wide, upturned subgenital plate (Figs 147 , 161 ) lacking spines, pre-apically inserted with short, downward pointing styli (Fig. 175 ) and in the narrow, completely fused slightly asymmetrical apical arms of the titillator (Figs 190 , 206 ). Females distinctly differ in the subgenital plate (Figs 46 , 60 ) with a central hump bordered by deep and wide semi-circular grooves. From all Cretan Eupholidoptera species, the black part in the anterior half of the pronotum is most pronounced in E. annamariae . For more details differentiating E. annamariae from other Cretan Eupholidoptera see Table 5 . Figures 69-82. Male anal tergite in dorsal view 69 Eupholidoptera smyrnensis Makrigiannis RMNH.INS152909 70 Eupholidoptera gemellata holotype Mt. Idhi RMNH.INS1105300 71 Eupholidoptera pallipes paratype Mt. Lefka RMNH.INS1105313 72 Eupholidoptera cretica Mt. Lefka, Omalos FC17807 RMNH.INS1141838 73 Eupholidoptera latens Mt. Lefka, Kalergi CT2003.009.01 74 Eupholidoptera giuliae Chora Sfakion CT2000.024.04 75 Eupholidoptera francisae sp. nov. holotype Andikithira CT2002.004.04 76 Eupholidoptera annamariae Kato Zakros CT2000.030.04 77 Eupholidoptera astyla Krotos RMNH.INS1141819 78 Eupholidoptera feri holotype Mt. Dikti, Katharo plain RMNH.INS 1105297 79 Eupholidoptera mariannae holotype Anatoli RMNH.INS1105311 80 Eupholidoptera forcipata Mt. Idhi CT1987.044.01 81 Eupholidoptera marietheresae sp. nov. paratype Mt. Dikti FC1606 CT2000.096.01 82 Eupholidoptera jacquelinae holotype Gavdos CT2000.005.02. Scale bars: 1 mm. Figures 83-96. Male anal tergite in caudal view 83 Eupholidoptera smyrnensis Makrigiannis RMNH.INS152909 84 Eupholidoptera gemellata Mt. Idhi CT2000.095.01 85 Eupholidoptera pallipes paratype Mt. Lefka RMNH.INS1105314 86 Eupholidoptera cretica Mt. Lefka, Omalos FC17807 RMNH.INS1141838 87 Eupholidoptera latens Mt. Lefka, Kalergi CT2003.009.01 88 Eupholidoptera giuliae Chora Sfakion CT2000.014.01 89 Eupholidoptera francisae sp. nov. paratype Andikithira CT2002.004.02 90 Eupholidoptera annamariae Kato Zakros CT2000.030.01 91 Eupholidoptera astyla Krotos RMNH.INS1141819 92 Eupholidoptera feri holotype Mt. Dikti, Katharo plain RMNH.INS1105297 93 Eupholidoptera mariannae Prina FC17798 RMNH.INS1141840 94 Eupholidoptera forcipata Mt. Idhi CT1987.044.01 95 Eupholidoptera marietheresae sp. nov. paratype Mt. Dikti FC1606 CT2000.096.01 96 Eupholidoptera jacquelinae paratype Gavdos NHMC2001.004.12. Scale bars: 1 mm. Figures 97-110. Male anal tergite in lateral view 97 Eupholidoptera smyrnensis Makrigiannis RMNH5087053 98 Eupholidoptera gemellata Mt. Idhi CT2000.095.01 99 Eupholidoptera pallipes paratype Mt. Lefka RMNH.INS1105314 100 Eupholidoptera cretica Mt. Lefka, Omalos FC17807 RMNH.INS1141838 101 Eupholidoptera latens Mt. Lefka, Kalergi CT1987.047.01 102 Eupholidoptera giuliae Argoules CT1995.011.11 103 Eupholidoptera francisae sp. nov. paratype Andikithira CT2002.004.02 104 Eupholidoptera annamariae Kato Zakros CT2000.030.03 105 Eupholidoptera astyla Ano Viannos CT1995.024.01 106 Eupholidoptera feri holotype Mt. Dikti, Katharo plain RMNH.INS1105297 107 Eupholidoptera mariannae Kalavros CT2017.029.01 108 Eupholidoptera forcipata Mt. Idhi CT1987.044.01 109 Eupholidoptera marietheresae sp. nov. paratype Mt. Dikti FC1606 CT2000.096.01 110 Eupholidoptera jacquelinae Gavdopoula CT1996.019.01. Scale bars: 1 mm. Figures 111-125. Male cercus in dorsal view 111 Eupholidoptera smyrnensis Makrigiannis RMNH5087053 112 Eupholidoptera gemellata Mt. Idhi FC1651 RMNH.INS1141843 113 Eupholidoptera pallipes Mt. Lefka paratype RMNH.INS1105314 114 Eupholidoptera cretica Mt. Lefka, Omalos FC17807 RMNH.INS1141838 115 Eupholidoptera latens Xiloskalo IBER DC-Ort000558 116 Eupholidoptera giuliae Chora Sfakion CT2000.014.02 117 Eupholidoptera francisae sp. nov. holotype Andikithira CT2002.004.04 118 Eupholidoptera annamariae Kato Zakros CT2000.030.03 119 Eupholidoptera annamariae Xerokampos, male with anomalous cercus, IBER DC-Ort000565 120 Eupholidoptera astyla Ierapetra MfN s.n. 121 Eupholidoptera feri holotype Mt. Dikti, Katharo plain RMNH.INS1105297 122 Eupholidoptera mariannae holotype Anatoli RMNH.INS1105311 123 Eupholidoptera forcipata Mt. Idhi CT1987.044.01 124 Eupholidoptera marietheresae sp. nov. holotype Mt. Dikti FC1606 RMNH.INS1141850 125 Eupholidoptera jacquelinae holotype Gavdos CT2000.005.02. Scale bars: 1 mm. Figures 126-139. Male cercus in lateral view 126 Eupholidoptera smyrnensis Makrigiannis RMNH.5087053 127 Eupholidoptera gemellata Mt. Idhi FC1651 RMNH.INS1141843 128 Eupholidoptera pallipes paratype Mt. Lefka RMNH.INS1105314 129 Eupholidoptera cretica Mt. Lefka, Omalos FC17807 RMNH.INS1141838 130 Eupholidoptera latens Mt. Lefka Kalergi CT2003.009.01 131 Eupholidoptera giuliae Chora Sfakion CT2000.024.04 132 Eupholidoptera francisae paratype sp. nov. Andikithira CT2002.004.08 133 Eupholidoptera annamariae Kato Zakros CT1995.020.02 134 Eupholidoptera astyla paratype Ierapetra MfN s.n. 135 Eupholidoptera feri holotype Mt. Dikti, Katharo plain RMNH.INS1105297 136 Eupholidoptera mariannae holotype Anatoli RMNH.INS1105311 137 Eupholidoptera forcipata Mt. Idhi CT1987.044.01 138 Eupholidoptera marietheresae sp. nov. holotype Mt. Dikti FC1606 RMNH.INS1141850 139 Eupholidoptera jacquelinae holotype Gavdos CT2000.005.02. Scale bars: 1 mm. Figures 140-153. Male subgenital plate in ventral view 140 Eupholidoptera smyrnensis Lagkada RMNH 5106270 141 Eupholidoptera gemellata Mt. Idhi FC1651 RMNH.INS1141843 142 Eupholidoptera pallipes paratype Mt. Lefka RMNH.INS1105313 143 Eupholidoptera cretica Mt. Lefka, Omalos FC17807 RMNH.INS1141838 144 Eupholidoptera latens Mt. Lefka Kalergi CT1987.047.02 145 Eupholidoptera giuliae Skaloti IBER DC-Ort000564 146 Eupholidoptera francisae sp. nov. holotype Andikithira CT2002.004.04 147 Eupholidoptera annamariae Kato Zakros CT2000.030.04 148 Eupholidoptera astyla Krotos RMNH.INS1141819 149 Eupholidoptera feri holotype Mt. Dikti, Katharo plain RMNH.INS1105297 150 Eupholidoptera mariannae holotype Anatoli RMNH.INS1105311 151 Eupholidoptera forcipata Mt. Idhi CT1987.044.03 152 Eupholidoptera marietheresae sp. nov. holotype Mt. Dikti FC1606 RMNH.INS1141850 153 Eupholidoptera jacquelinae holotype Gavdos 2000.005.02. Scale bars: 1 mm. Figures 154-167. Male subgenital plate in lateral view 154 Eupholidoptera smyrnensis Lagkada RMNH.INS152909 155 Eupholidoptera gemellata holotype Mt. Idhi RMNH.INS1105300 156 Eupholidoptera pallipes paratype Mt. Lefka RMNH.INS1105313 157 Eupholidoptera cretica Mt. Lefka, Omalos FC17807 (RMNH.INS1141838 158 Eupholidoptera latens Mt. Lefka Kalergi CT2003.009.01 159 Eupholidoptera giuliae Skaloti IBER DC-Ort000564 160 Eupholidoptera francisae sp. nov. holotype Andikithira CT2002.004.04 161 Eupholidoptera annamariae Kato Zakros CT2000.030.04 162 Eupholidoptera astyla Krotos RMNH5086992 163 Eupholidoptera feri holotype Mt. Dikti, Katharo plain RMNH.INS1105297 164 Eupholidoptera mariannae holotype Anatoli RMNH.INS1105311 165 Eupholidoptera forcipata Mt. Idhi CT1987.044.03 166 Eupholidoptera marietheresae sp. nov. holotype Mt. Dikti FC1606 RMNH.INS1141850 167 Eupholidoptera jacquelinae holotype Gavdos CT2000.005.02. Scale bars: 1 mm. Figures 168-181. Male stylus in lateral view 168 Eupholidoptera smyrnensis Makrigiannis RMNH.INS152909 169 Eupholidoptera gemellata holotype Mt. Idhi RMNH.INS1105300 170 Eupholidoptera pallipes paratype Mt. Lefka RMNH.INS1105313 171 Eupholidoptera cretica Mt. Lefka, Omalos FC17807 RMNH.INS1141838 172 Eupholidoptera latens Mt. Lefka Kalergi CT2003.009.01 173 Eupholidoptera giuliae Skaloti IBER DC-Ort000564 174 Eupholidoptera francisae sp. nov. paratype Andikithira CT2002.004.08 175 Eupholidoptera annamariae Kato Zakros CT2000.030.04 176 Eupholidoptera astyla Krotos RMNH.INS1141819 177 Eupholidoptera feri holotype Mt. Dikti, Katharo plain RMNH.INS1105297 178 Eupholidoptera mariannae holotype Anatoli RMNH.INS1105311 179 Eupholidoptera forcipata Mt. Idhi CT1987.044.03 180 Eupholidoptera marietheresae sp. nov. holotype Mt. Dikti FC1606 RMNH.INS1141850 181 Eupholidoptera jacquelinae holotype Gavdos CT2000.005.02. Scale bars: 1 mm. Figures 182-197. Male titillator in ventral view 182 Eupholidoptera smyrnensis Makrigiannis RMNH5087053 183 Eupholidoptera gemellata Mt. Idhi Amariou FC1651 RMNH.INS1141843 184 Eupholidoptera pallipes paratype Mt. Lefka RMNH.INS1105314 185 Eupholidoptera cretica Mt. Lefka, Omalos FC17807 RMNH.INS1141838 186 Eupholidoptera latens Mt. Lefka Kalergi CT2003.009.01 187 Eupholidoptera latens Rhodopos CH 8236 188 Eupholidoptera giuliae Argoules CT1995.011.11 189 Eupholidoptera francisae sp. nov. paratype Andikithira CT2002.004.08 190 Eupholidoptera annamariae Kato Zakros CT2000.030.01 191 Eupholidoptera astyla Krotos RMNH.INS1141819 192 Eupholidoptera astyla Kofinas FC460 RMNH.INS1141836 193 Eupholidoptera feri holotype Mt. Dikti, Katharo plain RMNH.INS1105297 194 Eupholidoptera mariannae paratype Malles CH2906A 195 Eupholidoptera forcipata Mt. Idhi CT2019.047.01 196 Eupholidoptera marietheresae sp. nov. paratype Mt. Dikti FC1606 CT2000.096.01 197 Eupholidoptera jacquelinae holotype Gavdos CT2000.005.02. Scale bars: 1 mm. Figures 198-212. Male titillator in lateral view 198 Eupholidoptera smyrnensis Makrigiannis RMNH5087053 199 Eupholidoptera gemellata Mt. Idhi Amariou FC1651 RMNH.INS1141843 200 Eupholidoptera pallipes paratype Mt. Lefka RMNH.INS1105314 201 Eupholidoptera cretica Mt. Lefka, Omalos FC17807 RMNH.INS1141838 202 Eupholidoptera latens Mt. Lefka Kalergi CT2003.009.01 203 Eupholidoptera latens Rhodopos CH 8236 204 Eupholidoptera giuliae Argoules CT1995.011.11 205 Eupholidoptera francisae sp. nov. paratype Andikithira CT2002.004.08 206 Eupholidoptera annamariae Kato Zakros CT2000.030.01 207 Eupholidoptera astyla Krotos RMNH.INS1141819 208 Eupholidoptera feri holotype Mt. Dikti, Katharo plain RMNH.INS1105297 209 Eupholidoptera mariannae Kalavros CT2017.029.01 210 Eupholidoptera forcipata Mt. Idhi CT1987.044.01 211 Eupholidoptera marietheresae sp. nov. holotype Mt. Dikti FC1606 RMNH.INS1141850 212 Eupholidoptera jacquelinae holotype Gavdos CT2000.005.02. Scale bars: 1 mm. Distribution. The species was described from Kato Zakros along the eastern coast of Crete. After its original finding it was collected again in the same area between Kato Zakros and Zakros ( Ciplak et al. 2009 ). New data presented here show that the range extends from the southeastern coast east of Kalo Nero Bay along the entire east coast up into the northernmost peninsular and along the northern coast westward up to and beyond Sitia (Fig. 254 ). It is still unclear if the species also occurs more inland east of the Koutsouras-Sitia road. For a complete list of localities, specimens and repositories see Suppl. material 1. Habitat. The species has been found in sparse phrygana between sea level and 550 m in dry open terrains with bare ground, covered with small spiny or thorny shrublets in which it hides during the day. The species was also found in a pitfall trap in sand dunes near Xerokampos along the southeastern coast. Phenology. Hand catches of this species were made between end of May and mid-August (25/05-15/08). This roughly coincides with the period during which the species was caught in pitfall traps. Still their presence may be more prolonged into August or up to October as a trap sampled 12 October 2000 and set 6 August still contained nine adults.