Eupholidoptera astyla (Ramme, 1927)

Willemse, Luc, Tilmans, Jos, Kotitsa, Nefeli, Trichas, Apostolos, Heller, Klaus-Gerhard, Chobanov, Dragan & Ode, Baudewijn, 2023, A review of Eupholidoptera (Orthoptera, Tettigoniidae) from Crete, Gavdos, Gavdopoula, and Andikithira, ZooKeys 1151, pp. 67-158 : 67

Eupholidoptera astyla (Ramme, 1927)

Figs 10 , 19 , 33 , 47 , 61 , 77 , 91 , 105 , 120 , 134 , 148 , 162 , 176 , 191 , 192 , 207 , 213 , 224 , 225 , 241 , 254 , 256 , 259

Pholidoptera astyla Ramme 1927: 133.

Eupholidoptera astyla (Ramme, 1927); Ramme 1951: 198.

Eupholidoptera astyla Morphological description. Ramme 1927: 133; 1939: 100; Harz 1969: 377.

Eupholidoptera astyla Bioacoustics. Çiplak et al. 2009: 27, 54-55.

Examined specimens.

1 ♂ (paratype); 81 ♂, 59 ♀ (for details see Suppl. material 2).

Diagnostic features.

Frontal part of head (Fig. 19 ) pale with black dots; frontal half of pronotal disc (Fig. 33 ) with more or less extensive black patch, border with pale rear half transverse or V-shaped. Male (Fig. 241 ) - stridulatory file with 101-105 teeth (including proximal and distal ones), density of teeth in middle two thirds of the file 22-24 teeth per mm; anal tergite (Figs 77 , 91 , 105 ) with hind margin toward middle forming two small pointed teeth separated by a short narrow V-shaped excision, tips pointing downwards; cerci (Figs 120 , 134 ) unarmed, 5-7 × longer than wide, weakly conical, weakly curved inward in basal half, in profile straight; subgenital plate (Figs 148 , 162 ) wider than long, widest halfway, sides rimmed, in profile upturned, tip apical lobes narrowly truncate, spineless, at inner side emarginate with V-shaped excision along one fifth of length; styli (Fig. 176 ) minute, circular, inserted ventrally, proximal of tip of apical lobe, pointing inward to downward; titillator (Figs 191 , 207 ) asymmetrical, apical arms widening from base, halfway splitting into two strongly thickened, flattened and wrinkled arms, forming wide to almost straight angle ending into two very strong curved spine-like teeth, pointing left or right.

Redescription of female.

In 1927 E. astyla was described based on a single male from Naxos and three females and the male abdomen from Crete ( Ramme 1927). According to Willemse and Heller (2001) the presence of E. astyla on Naxos is doubtful. An extensive search on Naxos in June 2019 by the first author only revealed E. smyrnensis at midlevel altitudes in the north-eastern part of the island but no other Eupholidoptera species. Willemse and Heller (2001) indicated that the E. astyla females listed in the original description ( Ramme 1927: 134) from Ierapetra, Anatoli, and Kato Chorio could in fact belong to E. mariannae which they described in their paper from this area. In 2001 female E. mariannae were not known and this assumption could neither be confirmed nor denied. Based on the discovery of male and female E. mariannae in neighbouring locations near Kavousi and Kalavros, opportunity has been taken to re-examine paratypes used by Ramme (1927) in his description of E. astyla and compare them with female E. astyla and E. mariannae described in this paper. Unfortunately, the female from “Jerapetra” could not be located in the MfNB. Comparison of the subgenital plates (Figs 214 , 215 ) revealed that females from Kato Chorio and Anadoli resemble female E. mariannae from Kalavros and Kavousi rather than E. astyla . This implies that Ramme based his description of female E. astyla on female E. mariannae . Consequently female E. astyla has still not been described and is redescribed here based on specimens from Rethimno and Iraklion.

Description.

Female.

Examined specimens.

11 ♀: RETHIMNO: Idhi Mt., Idhaio Andro -1987.041.02 (CT); Idhi Mt., Ski-centre - 1987.046.03 (CT); Nea Kria Vrisi, 2 km NW - 2000.016.04 (CT); IRAKLION: Ano Viannos, 3 km SE - 1995.024.03 (CT), RMNH.5014909 (RMNH), RMNH.5086970 (RMNH), RMNH 5086971 (RMNH); Krotos, 0.5 km N - RMNH.INS1141820 (RMNH); Marathos - RMNH.5086989 (RMNH); Mournia, 3 km SW - 2001.008.04 (CT); Tsoutsouros, 6 km NNW - 1995.014.03 (CT). For more details see Suppl. material 2.

General appearance (Figs 224 , 225 ) and colouration as male. First abdominal segment dorsally black, laterally lighter; remaining segments dorsally yellowish brown, sides lighter coloured, last segment completely black. Elytra in dorsal view covered by pronotum, in profile barely protruding, light coloured. Cercus short, conical tapering, more so in apical third toward a pointed tip, straight to slightly upturned in profile, straight to slightly curved inward in dorsal view, covered with pale short and long hairs. Subgenital plate (Figs 47 , 61 ) oblong, greatest width halfway or in distal quarter, in ventral view convex, proximally flattened to slight depressed, halfway on side with or without a (in-)distinct bulge, surface shiny, smooth with dispersed hairs, hind margin converging gradually from halfway or more abruptly in distal quarter, with medial V-shaped excision along one third of length, corners rectangular to sharp-angled, in profile triangular to trapezoid with a more or less distinct dorsal depression, lower edge straight distally slightly upturned, tip obtuse angular. Ovipositor almost straight to slightly upcurved, 1.5-2.0 × longer than pronotum.

Measurements.

See Tables 6 , 7 .

Bioacoustics.

Based upon the sound recordings of two specimens (20 syllables measured), the song of E. astyla , 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. astyla , the syllable duration is ~ 120 ms, with a syllable rate up to ~ 1/s. Published records ( Çiplak et al. 2009) show a syllable duration of ~ 166 ms at 25 °C and a syllable repetition rate of less than 1/s. 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 astyla see Suppl. material 3.

Variation.

High altitude specimens are smaller than specimens found at lower altitudes. Variation in black and pale colour patterns seem linked to individuals rather than to populations. Cercus more or less slender and more or less bent inward. Medial excision anal tergite V- to U-shaped, adjoining teeth aligned with dorsal surface pointing distally or bent downward, pointing downward. Subgenital plate more or less compact, in profile, lower margin evenly rounded or with an angle halfway. Styli minute to small, in the west and north pointing inward, toward the south and east pointing downward, exceptionally also outward. In some males from Asterousia Mt. (central-south Crete), styli were lacking almost completely. Titillator can be more or less compact, apical arms parallel or slightly divergent, apical teeth gradually or suddenly and more strongly pointed, pointing right or left. Two males (of 34) collected in pitfall traps near Kofinas along the south coast showed an almost symmetrical titillator, the two apical arms pointing in opposite directions (Fig. 192 ). In other characters these two males fully matched E. astyla , as did all other males from Kofinas. The symmetrical titillators are considered individual anomalies. The shape of the female subgenital plate varies: in the west and at higher altitudes on Mts. Idi and Dikti being plump with a short median excision, toward the east changing to oblong more distinctly acutely bilobed with a deeper excision, resembling those of E. feri and E. mariannae .

Differential diagnosis.

Males differ from congenerics in the strongly asymmetrical, thickened and wrinkled apical arms of the titillator (Figs 191 , 207 ) pointing left or right, in the narrow V-shaped excision in the anal tergite (Figs 77 , 91 , 105 ) with tips pointing downward, in the slender, unarmed weakly inward curved cerci (Figs 120 , 134 ), in the wide, upturned, spineless subgenital plate (Figs 148 , 162 ) and in the minute, pre-apically inserted styli pointing downward (Fig. 176 ). Females differ in the oblong subgenital plate (Figs 47 , 61 ), proximally convex with a short excision. In the southeast toward Lasithi female subgenital plates are longer, the apical lobes on both sides of the medial excision more pointed, resembling E. mariannae . In colouration, particularly the anterior half of the pronotum E. astyla resembles E. annamariae , E. feri , E. giuliae , and E. mariannae . For more details differentiating E. astyla from other Cretan Eupholidoptera see Table 5 .

Distribution.

From the Cretan species of Eupholidoptera , E. astyla has the widest range. Current data indicate its range covers large parts of central Crete, stretching from central and eastern Rethimno to western Lasithi, from Skaleta east of the city of Rethimno in the northwest to Ierapetra along the southern coast in the southeast (Fig. 254 ). Despite the additional localities presented here, the exact boundaries in the west, where it meets E. giuliae , and in the east, where it meets E. mariannae and E. feri , are not clear. Current information suggests E. astyla to be absent from large parts of the central lowlands south of Iraklion and the northern coastal region east of Iraklion but this certainly requires additional investigation. Opportunity was taken to examine (and photograph) the male mentioned by Ramme (1927) in his description of E. astyla of which only the last part of the abdomen was left. Based on the shape of the titillator (Fig. 213 ) this male clearly belongs to E. astyla . The label of this male only states “Jerapetra”. It is not clear whether the name “Jerapetra” actually refers to the town Ierapetra or the district. Either way, assuming the label is correct, the presence in Ierapetra indicates there is an area west of Ierapetra where both E. astyla and E. mariannae may occur together (Fig. 254 ). For a complete list of localities, specimens and repositories see Suppl. material 1.

Habitat.

The species habitats cover a wide altitudinal range: from sea level along the northern and southern coast to 1550-1800 m on Mt. Idi and Mt. Dikti. Pitfall traps that caught E. astyla were placed in sparse to dense phrygana, maquis and areas dominated by pine trees.

Phenology.

Hand catches indicate adults can be found from early May onward at lower altitudes up to the end of August at higher altitudes. Pitfall trap catches indicate that especially at higher altitudes the species can be found at least up to the second half of September or even early October.

Figures

Figure 256. Bayesian inference phylogenetic tree of Eupholidoptera of Crete, Andikithira and Gavdos, including two outgroups and congenerics from the Greek and Turkish mainlands, based on a 1684 bp concatenated alignment of the NADH 2 + ITS fragments. Node values at branches show node support with BI posterior probabilities. Different colours correspond to the different morphological species. See Suppl. material 4 for details of the specimens and Suppl. material 5 for details of NADH 2 and IT IS sequences.

Figure 259. Mean syllable duration in recorded species from Crete (incl. Gavdos and Andikithira). See Suppl. material 3 for measurements per recorded specimen.

Figures 8 - 10. Colour pattern abdomen from above 8 Eupholidoptera smyrnensis ♀ Makrigiannis RMNH. INS 152908 9 Eupholidoptera gemellata ♀ Mt. Idhi FC 1651 RMNH. INS 1141844 10 Eupholidoptera astyla ♂ Krotos RMNH 5086990.

Figures 11 - 24. Colour pattern of head, frontal view 11 Eupholidoptera smyrnensis ♀ Makrigiannis RMNH 5014918 12 Eupholidoptera gemellata ♂ Mt. Idhi RMNH. INS 1141843 13 Eupholidoptera pallipes ♂ paratype Mt. Lefka RMNH. INS 1105313 14 Eupholidoptera cretica ♂ Mt. Lefka, Omalos RMNH. INS 114838 15 Eupholidoptera latens ♀ Mt. Lefka, Kalergi CT 1987.047.04 16 Eupholidoptera giuliae ♂ Argoules CT 1995.011.18 17 Eupholidoptera francisae sp. nov. ♀ allotype Andikithira CT 2002.004.11 18 Eupholidoptera annamariae ♀ Kato Zakros CT 1995.020.03 19 Eupholidoptera astyla ♀ Ano Viannos CT 1987.024.03 20 Eupholidoptera feri ♂ holotype Mt. Dikti, Katharo plain RMNH. INS 1105297 21 Eupholidoptera mariannae ♀ Ag. Ioannis RMNH. 5014906 22 Eupholidoptera forcipata ♂ Mt. Idhi CT 1987.044.03 23 Eupholidoptera marietheresae sp. nov. ♂ paratype Mt. Dikti, FC 1606 CT 2000.096.01 24 Eupholidoptera jacquelinae ♂ holotype Gavdos CT 2000.005.02. Scale bars: 5 mm.

Figures 25 - 38. Colour pattern of pronotum, dorsal view 25 Eupholidoptera smyrnensis ♂ Makrigiannis RMNH. INS 152909 26 Eupholidoptera gemellata ♂ Mt. Idhi, Amariou RMNH. INS 1141843 27 Eupholidoptera pallipes ♂ paratype Mt. Lefka RMNH. INS 1105313 28 Eupholidoptera cretica ♂ Mt. Lefka, Omalos RMNH. INS 1141838 29 Eupholidoptera latens ♂ Rhodopos CH 8236 30 Eupholidoptera giuliae Chora Sfakion ♂ CT 2000.014.10 31 Eupholidoptera francisae sp. nov. ♂ holotype Andikithira CT 2002.004.04 32 Eupholidoptera annamariae ♂ Kato Zakros CT 2000.030.04 33 Eupholidoptera astyla ♂ Krotos RMNH. INS 1141819 34 Eupholidoptera feri ♂ holotype Mt. Dikti, Katharo plain RMNH. INS 1105297 35 Eupholidoptera mariannae ♀ Kalavros RMNH. 5086974 36 Eupholidoptera forcipata ♂ Mt. Idhi CT 1987.044.03 37 Eupholidoptera marietheresae sp. nov. ♂ holotype Mt. Dikti RMNH. INS 1141850 38 Eupholidoptera jacquelinae ♂ holotype Gavdos CT 2000.005.02. Scale bars: 5 mm.

Figures 39 - 52. Female subgenital plate in ventral view 39 Eupholidoptera smyrnensis Makrigiannis RMNH. 5014918 40 Eupholidoptera gemellata Mt. Idhi, FC 1602 CT 2000.095.02 41 Eupholidoptera pallipes allotype Mt. Lefka RMNH. INS 1105312 42 Eupholidoptera cretica Mt. Lefka, Omalos RMNH. INS 1141837 43 Eupholidoptera latens Mt. Lefka, Kalergi CT 1987.047.04 44 Eupholidoptera giuliae Argoules CT 1995.011.03 45 Eupholidoptera francisae sp. nov. allotype Andikithira CT 2002.004.11 46 Eupholidoptera annamariae Kato Zakros CT 1995.020.03 47 Eupholidoptera astyla Krotos RMNH. INS 1141820 48 Eupholidoptera feri allotype Mt. Dikti, Katharo plain RMNH. INS 1105298 49 Eupholidoptera mariannae Kalavros RMNH 5014912 50 Eupholidoptera forcipata Mt. Idhi, FC 1602 RMNH. INS 1141845 51 Eupholidoptera marietheresae sp. nov. allotype Mt. Dikti RMNH. INS 1141849 52 Eupholidoptera jacquelinae Gavdos allotype CT 2001.004.13. Scale bars: 1 mm.

Figures 53 - 66. Female subgenital plate in lateral view 53 Eupholidoptera smyrnensis Makrigiannis RMNH. 5014918 54 Eupholidoptera gemellata Mt. Idhi, FC 1602 CT 2000.095.02 55 Eupholidoptera pallipes allotype Mt. Lefka RMNH. INS 1105312 56 Eupholidoptera cretica Mt. Lefka, Omalos RMNH. INS 1141837 57 Eupholidoptera latens Mt. Lefka, Kalergi CT 1987.047.04 58 Eupholidoptera giuliae Argoules CT 1995.011.03 59 Eupholidoptera francisae sp. nov. allotype Andikithira CT 2002.004.11 60 Eupholidoptera annamariae Kato Zakros CT 1995.020.03 61 Eupholidoptera astyla Krotos RMNH. INS 1141820 62 Eupholidoptera feri allotype Mt. Dikti, Katharo plain RMNH. INS 1105298 63 Eupholidoptera mariannae Kalavros RMNH 5014912 64 Eupholidoptera forcipata Mt. Idhi, FC 1602 RMNH. INS 1141845 65 Eupholidoptera marietheresae sp. nov. allotype Mt. Dikti RMNH. INS 1141849 66 Eupholidoptera jacquelinae allotype Gavdos CT 2001.004.13. Scale bars: 1 mm.

Figures 69 - 82. Male anal tergite in dorsal view 69 Eupholidoptera smyrnensis Makrigiannis RMNH. INS 152909 70 Eupholidoptera gemellata holotype Mt. Idhi RMNH. INS 1105300 71 Eupholidoptera pallipes paratype Mt. Lefka RMNH. INS 1105313 72 Eupholidoptera cretica Mt. Lefka, Omalos FC 17807 RMNH. INS 1141838 73 Eupholidoptera latens Mt. Lefka, Kalergi CT 2003.009.01 74 Eupholidoptera giuliae Chora Sfakion CT 2000.024.04 75 Eupholidoptera francisae sp. nov. holotype Andikithira CT 2002.004.04 76 Eupholidoptera annamariae Kato Zakros CT 2000.030.04 77 Eupholidoptera astyla Krotos RMNH. INS 1141819 78 Eupholidoptera feri holotype Mt. Dikti, Katharo plain RMNH. INS 1105297 79 Eupholidoptera mariannae holotype Anatoli RMNH. INS 1105311 80 Eupholidoptera forcipata Mt. Idhi CT 1987.044.01 81 Eupholidoptera marietheresae sp. nov. paratype Mt. Dikti FC 1606 CT 2000.096.01 82 Eupholidoptera jacquelinae holotype Gavdos CT 2000.005.02. Scale bars: 1 mm.

Figures 83 - 96. Male anal tergite in caudal view 83 Eupholidoptera smyrnensis Makrigiannis RMNH. INS 152909 84 Eupholidoptera gemellata Mt. Idhi CT 2000.095.01 85 Eupholidoptera pallipes paratype Mt. Lefka RMNH. INS 1105314 86 Eupholidoptera cretica Mt. Lefka, Omalos FC 17807 RMNH. INS 1141838 87 Eupholidoptera latens Mt. Lefka, Kalergi CT 2003.009.01 88 Eupholidoptera giuliae Chora Sfakion CT 2000.014.01 89 Eupholidoptera francisae sp. nov. paratype Andikithira CT 2002.004.02 90 Eupholidoptera annamariae Kato Zakros CT 2000.030.01 91 Eupholidoptera astyla Krotos RMNH. INS 1141819 92 Eupholidoptera feri holotype Mt. Dikti, Katharo plain RMNH. INS 1105297 93 Eupholidoptera mariannae Prina FC 17798 RMNH. INS 1141840 94 Eupholidoptera forcipata Mt. Idhi CT 1987.044.01 95 Eupholidoptera marietheresae sp. nov. paratype Mt. Dikti FC 1606 CT 2000.096.01 96 Eupholidoptera jacquelinae paratype Gavdos NHMC 2001.004.12. Scale bars: 1 mm.

Figures 97 - 110. Male anal tergite in lateral view 97 Eupholidoptera smyrnensis Makrigiannis RMNH 5087053 98 Eupholidoptera gemellata Mt. Idhi CT 2000.095.01 99 Eupholidoptera pallipes paratype Mt. Lefka RMNH. INS 1105314 100 Eupholidoptera cretica Mt. Lefka, Omalos FC 17807 RMNH. INS 1141838 101 Eupholidoptera latens Mt. Lefka, Kalergi CT 1987.047.01 102 Eupholidoptera giuliae Argoules CT 1995.011.11 103 Eupholidoptera francisae sp. nov. paratype Andikithira CT 2002.004.02 104 Eupholidoptera annamariae Kato Zakros CT 2000.030.03 105 Eupholidoptera astyla Ano Viannos CT 1995.024.01 106 Eupholidoptera feri holotype Mt. Dikti, Katharo plain RMNH. INS 1105297 107 Eupholidoptera mariannae Kalavros CT 2017.029.01 108 Eupholidoptera forcipata Mt. Idhi CT 1987.044.01 109 Eupholidoptera marietheresae sp. nov. paratype Mt. Dikti FC 1606 CT 2000.096.01 110 Eupholidoptera jacquelinae Gavdopoula CT 1996.019.01. Scale bars: 1 mm.

Figures 111 - 125. Male cercus in dorsal view 111 Eupholidoptera smyrnensis Makrigiannis RMNH 5087053 112 Eupholidoptera gemellata Mt. Idhi FC 1651 RMNH. INS 1141843 113 Eupholidoptera pallipes Mt. Lefka paratype RMNH. INS 1105314 114 Eupholidoptera cretica Mt. Lefka, Omalos FC 17807 RMNH. INS 1141838 115 Eupholidoptera latens Xiloskalo IBER DC-Ort 000558 116 Eupholidoptera giuliae Chora Sfakion CT 2000.014.02 117 Eupholidoptera francisae sp. nov. holotype Andikithira CT 2002.004.04 118 Eupholidoptera annamariae Kato Zakros CT 2000.030.03 119 Eupholidoptera annamariae Xerokampos, male with anomalous cercus, IBER DC-Ort 000565 120 Eupholidoptera astyla Ierapetra MfN s. n. 121 Eupholidoptera feri holotype Mt. Dikti, Katharo plain RMNH. INS 1105297 122 Eupholidoptera mariannae holotype Anatoli RMNH. INS 1105311 123 Eupholidoptera forcipata Mt. Idhi CT 1987.044.01 124 Eupholidoptera marietheresae sp. nov. holotype Mt. Dikti FC 1606 RMNH. INS 1141850 125 Eupholidoptera jacquelinae holotype Gavdos CT 2000.005.02. Scale bars: 1 mm.

Figures 126 - 139. Male cercus in lateral view 126 Eupholidoptera smyrnensis Makrigiannis RMNH. 5087053 127 Eupholidoptera gemellata Mt. Idhi FC 1651 RMNH. INS 1141843 128 Eupholidoptera pallipes paratype Mt. Lefka RMNH. INS 1105314 129 Eupholidoptera cretica Mt. Lefka, Omalos FC 17807 RMNH. INS 1141838 130 Eupholidoptera latens Mt. Lefka Kalergi CT 2003.009.01 131 Eupholidoptera giuliae Chora Sfakion CT 2000.024.04 132 Eupholidoptera francisae paratype sp. nov. Andikithira CT 2002.004.08 133 Eupholidoptera annamariae Kato Zakros CT 1995.020.02 134 Eupholidoptera astyla paratype Ierapetra MfN s. n. 135 Eupholidoptera feri holotype Mt. Dikti, Katharo plain RMNH. INS 1105297 136 Eupholidoptera mariannae holotype Anatoli RMNH. INS 1105311 137 Eupholidoptera forcipata Mt. Idhi CT 1987.044.01 138 Eupholidoptera marietheresae sp. nov. holotype Mt. Dikti FC 1606 RMNH. INS 1141850 139 Eupholidoptera jacquelinae holotype Gavdos CT 2000.005.02. Scale bars: 1 mm.

Figures 140 - 153. Male subgenital plate in ventral view 140 Eupholidoptera smyrnensis Lagkada RMNH 5106270 141 Eupholidoptera gemellata Mt. Idhi FC 1651 RMNH. INS 1141843 142 Eupholidoptera pallipes paratype Mt. Lefka RMNH. INS 1105313 143 Eupholidoptera cretica Mt. Lefka, Omalos FC 17807 RMNH. INS 1141838 144 Eupholidoptera latens Mt. Lefka Kalergi CT 1987.047.02 145 Eupholidoptera giuliae Skaloti IBER DC-Ort 000564 146 Eupholidoptera francisae sp. nov. holotype Andikithira CT 2002.004.04 147 Eupholidoptera annamariae Kato Zakros CT 2000.030.04 148 Eupholidoptera astyla Krotos RMNH. INS 1141819 149 Eupholidoptera feri holotype Mt. Dikti, Katharo plain RMNH. INS 1105297 150 Eupholidoptera mariannae holotype Anatoli RMNH. INS 1105311 151 Eupholidoptera forcipata Mt. Idhi CT 1987.044.03 152 Eupholidoptera marietheresae sp. nov. holotype Mt. Dikti FC 1606 RMNH. INS 1141850 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. INS 152909 155 Eupholidoptera gemellata holotype Mt. Idhi RMNH. INS 1105300 156 Eupholidoptera pallipes paratype Mt. Lefka RMNH. INS 1105313 157 Eupholidoptera cretica Mt. Lefka, Omalos FC 17807 (RMNH. INS 1141838 158 Eupholidoptera latens Mt. Lefka Kalergi CT 2003.009.01 159 Eupholidoptera giuliae Skaloti IBER DC-Ort 000564 160 Eupholidoptera francisae sp. nov. holotype Andikithira CT 2002.004.04 161 Eupholidoptera annamariae Kato Zakros CT 2000.030.04 162 Eupholidoptera astyla Krotos RMNH 5086992 163 Eupholidoptera feri holotype Mt. Dikti, Katharo plain RMNH. INS 1105297 164 Eupholidoptera mariannae holotype Anatoli RMNH. INS 1105311 165 Eupholidoptera forcipata Mt. Idhi CT 1987.044.03 166 Eupholidoptera marietheresae sp. nov. holotype Mt. Dikti FC 1606 RMNH. INS 1141850 167 Eupholidoptera jacquelinae holotype Gavdos CT 2000.005.02. Scale bars: 1 mm.

Figures 168 - 181. Male stylus in lateral view 168 Eupholidoptera smyrnensis Makrigiannis RMNH. INS 152909 169 Eupholidoptera gemellata holotype Mt. Idhi RMNH. INS 1105300 170 Eupholidoptera pallipes paratype Mt. Lefka RMNH. INS 1105313 171 Eupholidoptera cretica Mt. Lefka, Omalos FC 17807 RMNH. INS 1141838 172 Eupholidoptera latens Mt. Lefka Kalergi CT 2003.009.01 173 Eupholidoptera giuliae Skaloti IBER DC-Ort 000564 174 Eupholidoptera francisae sp. nov. paratype Andikithira CT 2002.004.08 175 Eupholidoptera annamariae Kato Zakros CT 2000.030.04 176 Eupholidoptera astyla Krotos RMNH. INS 1141819 177 Eupholidoptera feri holotype Mt. Dikti, Katharo plain RMNH. INS 1105297 178 Eupholidoptera mariannae holotype Anatoli RMNH. INS 1105311 179 Eupholidoptera forcipata Mt. Idhi CT 1987.044.03 180 Eupholidoptera marietheresae sp. nov. holotype Mt. Dikti FC 1606 RMNH. INS 1141850 181 Eupholidoptera jacquelinae holotype Gavdos CT 2000.005.02. Scale bars: 1 mm.

Figures 182 - 197. Male titillator in ventral view 182 Eupholidoptera smyrnensis Makrigiannis RMNH 5087053 183 Eupholidoptera gemellata Mt. Idhi Amariou FC 1651 RMNH. INS 1141843 184 Eupholidoptera pallipes paratype Mt. Lefka RMNH. INS 1105314 185 Eupholidoptera cretica Mt. Lefka, Omalos FC 17807 RMNH. INS 1141838 186 Eupholidoptera latens Mt. Lefka Kalergi CT 2003.009.01 187 Eupholidoptera latens Rhodopos CH 8236 188 Eupholidoptera giuliae Argoules CT 1995.011.11 189 Eupholidoptera francisae sp. nov. paratype Andikithira CT 2002.004.08 190 Eupholidoptera annamariae Kato Zakros CT 2000.030.01 191 Eupholidoptera astyla Krotos RMNH. INS 1141819 192 Eupholidoptera astyla Kofinas FC 460 RMNH. INS 1141836 193 Eupholidoptera feri holotype Mt. Dikti, Katharo plain RMNH. INS 1105297 194 Eupholidoptera mariannae paratype Malles CH 2906 A 195 Eupholidoptera forcipata Mt. Idhi CT 2019.047.01 196 Eupholidoptera marietheresae sp. nov. paratype Mt. Dikti FC 1606 CT 2000.096.01 197 Eupholidoptera jacquelinae holotype Gavdos CT 2000.005.02. Scale bars: 1 mm.

Figures 198 - 212. Male titillator in lateral view 198 Eupholidoptera smyrnensis Makrigiannis RMNH 5087053 199 Eupholidoptera gemellata Mt. Idhi Amariou FC 1651 RMNH. INS 1141843 200 Eupholidoptera pallipes paratype Mt. Lefka RMNH. INS 1105314 201 Eupholidoptera cretica Mt. Lefka, Omalos FC 17807 RMNH. INS 1141838 202 Eupholidoptera latens Mt. Lefka Kalergi CT 2003.009.01 203 Eupholidoptera latens Rhodopos CH 8236 204 Eupholidoptera giuliae Argoules CT 1995.011.11 205 Eupholidoptera francisae sp. nov. paratype Andikithira CT 2002.004.08 206 Eupholidoptera annamariae Kato Zakros CT 2000.030.01 207 Eupholidoptera astyla Krotos RMNH. INS 1141819 208 Eupholidoptera feri holotype Mt. Dikti, Katharo plain RMNH. INS 1105297 209 Eupholidoptera mariannae Kalavros CT 2017.029.01 210 Eupholidoptera forcipata Mt. Idhi CT 1987.044.01 211 Eupholidoptera marietheresae sp. nov. holotype Mt. Dikti FC 1606 RMNH. INS 1141850 212 Eupholidoptera jacquelinae holotype Gavdos CT 2000.005.02. Scale bars: 1 mm.

Figures 213 - 215. Eupholidoptera astyla paratypes 213 titillator in dorsal view Eupholidoptera astyla ♂ paratype Ierapetra s. n. 214 subgenital plate in ventral view Eupholidoptera mariannae ♀ Anadoli s. n. [paratype of Eupholidoptera astyla] 215 Eupholidoptera mariannae ♀ Kato Chorion s. n. [paratype of Eupholidoptera astyla].

Figures 224 - 235. Habitus Eupholidoptera spp. in dorsal and lateral view 224, 225 Eupholidoptera astyla ♀ Mt. Idhi CT 1987.046.03 226, 227 Eupholidoptera cretica ♀ Mt. Lefka Omalos FC 17807 RMNH. INS 1141837 228, 229 Eupholidoptera francisae sp. nov. ♂ holotype Andikithira CT 2002.004.04 230, 231 Eupholidoptera francisae sp. nov. ♀ allotype Andikithira CT 2002.004.11 232, 233 Eupholidoptera gemellata ♀ Mt. Idhi FC 1651 RMNH. INS 1141844 234, 235 Eupholidoptera mariannae ♀ Ag. Ioannis RMNH 5014906 1844. Scale bars: 10 mm.

Figures 240 - 246. Field images Eupholidoptera spp. Crete 240 Eupholidoptera annamariae ♂ Kata Zakros CT 1995.0530 241 Eupholidoptera astyla ♂ Mt. Idhi CT 2004.08.14 242 Eupholidoptera jacquelinae ♀ Gavdos 243 Eupholidoptera francisae sp. nov. ♂ paratype Andikithira CT 2002.004.07 244 Eupholidoptera francisae sp. nov. ♀ paratype Andikithira CT 2002.004.12 245 Eupholidoptera francisae ♂ paratype Marouliana RMNH 5087052 246 Eupholidoptera francisae ♀ paratype Marouliana RMNH 5014917.

Figures 254, 255. Distribution maps Eupholidoptera Crete and adjacent islands 254 lowland species 255 highland species.