Eupholidoptera francisae Tilmans & Ode, 2023

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 francisae Tilmans & Ode sp. nov.

Figs 17 , 31 , 45 , 59 , 67 , 75 , 89 , 103 , 117 , 132 , 146 , 160 , 174 , 189 , 205 , 216-217 , 218 , 228-231 , 243-246 , 254 , 256 , 257 , 258 , 259

Remark.

The Eupholidoptera populations present on the island of Andikithira and in the area of western-southwestern Chania in Crete not only proved to differ from the geographically nearest other taxa of the genus: Eupholidoptera spinigera , restricted to the island of Kithira, and Eupholidoptera latens from northern and central Chania, but also from all its other congenerics. This new taxon is described below. For arguments to assign the Eupholidoptera populations of Andikithira and western/southwestern Chania populations as one single new taxon see under Discussion.

Examined specimens.

Type specimens. ♂ holotype (2002.004.04) (CT), ♀ allotype (2002.004.11) (CT), both labeled: HELLAS, Andikithira, 150 m, 9.V.2002/3 km S.E.S. Potamos/WGS 84 35°51.996'N, 023°18.114'E /legnt. J.M. Tilmans and J.F.R. Tilmans-Smid.

Paratypes. 8 ♂ & 5 ♀ (CT), 1 ♂ & 1 ♀ (NHMC), 1 ♂ & 1 ♀ (RMNH): same location and date as holotype; further paratypes 2 ♀ (CT): HELLAS, Andikithira, 50 m, 9.V.2002/0,6 km S.E.S. Potamos/WGS84 35°52.600'N, 023°17.426'E /legnt. J.M. Tilmans & J.F.R. Tilmans-Smid; 1 ♂ & 2 ♀ (CT): HELLAS, Andikithira, 50 m, 27.V.2008/0,6 km S.E.S. Potamos/WGS84 35°52.605'N, 023°17.439'E /legnt. J.M. Tilmans & J.F.R. Tilmans-Smid; 1♂ & 2♀ (RMNH): Greece - Crete (Chania): Ag. Paraskevi (Elafonisos-Maniatiana)/445 m; 17.VI.2019; 35.285645°N, 23.588774°E /leg. L. Willemse & J. Tilmans; 1♀ (RMNH): Greece - Crete (Chania): 1 km NE of Anidhroi/385 m; 21.VI.2017; 35.255925°N, 23.737376°E /leg. L. Willemse & P. Zacharopoulou; 6♂ & 2♀ (CT): HELLAS, nomos Khania, 300 m/3 km E. Anidhroi, 27-29.IV.2001/ 35°15.288'N, 23°44.157'E /leg. J.M. Tilmans & J.F.R. Tilmans-Smid; 1♂ (CT), 2♂ & 1♀ (RMNH): Greece - Crete (Chania): 1 km N of Chondros/485 m; 16.VI.2019; 35.322094°N, 23.685799°E /leg L. Willemse & J. Tilmans; 1♂ & 1♀ (RMNH): Greece - Crete (Chania): Elos/480 m; 16.VI.2019; 35.367374°N, 23.637676°E)/leg. L. Willemse & J. Tilmans; 1♀ (CT), 1♀ (RMNH): Greece - Crete (Chania): 0.5 km W of Kamaria/345 m; 18.VI.2019; 35.282516°N, 23.778568°E /leg. L. Willemse & J. Tilmans; 1♀ (RMNH): 1 km S of Livadas/225 m; 18.VI.2019; 35.263004°N, 23.814722°E /leg. L. Willemse & J. Tilmans; 1♂ (IBER): Louchio, 0.5 km (35.3691°N, 23.6244°E) 665 m, 23/05/2018 Chobanov, D., Iorgu, I. & Borissov, S. 1♂ IBER; 1♂ 2♀ (RMNH): Greece - Crete (Chania); Marouliana, Ano Sfinari - Kostogiannides/715 m; 19.VI.2017; 35.390335°N, 23.605317°E /leg. L. Willemse & P. Zacharopoulou; 2♂ & 2♀ (CT), 3♂ & 4♀ (RMNH): Greece - Crete (Chania): Psariana-Aligi/420 m; 17.VI.2019; 35.351833°N, 23.694208°E /leg. L. Willemse & J. Tilmans; 1♂ & 1♀ (CT), 2♂ & 1♀ (RMNH): Greece - Crete (Chania): 1 km S of Sarakina/305 m; 16.VI.2019; 35.289181°N, 23.674417°E /leg. L. Willemse & J. Tilmans; 2♀ (IBER): Sfinari (35.4407°N, 23.5704°E) 1m, 23/05/2018 Chobanov, D., Iorgu, I. & Borissov, S.; 1♂ (CT), 1♂ & 1♀ (RMNH): Greece - Crete (Chania): just N of Strovles/420 m; 16.VI.2019; 35.368656°N, 23.669718°E /leg. L. Willemse & J. Tilmans; 1♂ (RMNH): Greece - Crete (Chania): 0.5 km N of Temenia/835 m; 17.VI.2019; 35.299652°N, 23.751684°E /leg. L. Willemse & J. Tilmans; 1♂ 1♀ (CT), 1♂ 2♀ (NHMC), 1♀ (RMNH): Greece - Crete (Rethimno): 1 km SE of Piso Moni Preveli/20 m; 12.VII.1997; 35.1518°N, 24.4725°E /leg. P. Lymberakis. (for details see Suppl. material 2).

Description.

Male. General appearance (Figs 228 , 229 ), elytra and legs as type species of genus, E. chabrieri .

Pronotum (Fig. 31 ) dorsally slightly flattened.

Forewing: stridulatory file left elytron consists of 96-138 teeth, shortest distance between proximal and distal end 3.0-3.9 mm, density of teeth in middle two thirds of the file 27-34 teeth per mm.

Anal tergite (Figs 75 , 89 , 103 ) apically strongly curved downward with round dorsomedian depression; posterior margin with wide, concave, moderately deep rounded (in many specimens semi-circular), median excision, bordered by two sharply toothed processes laterally, directed downward.

Cerci (Figs 117 , 132 ) long, slender, 6-7 × longer than greatest width, cylindrical with golden-coloured short and long hairs, without any tooth, slightly bent inwards.

Subgenital plate (Figs 146 , 160 ) very large, longer than wide, strikingly elongated, lateral margins swollen, ventrally with a median keel; basal third wide, then suddenly (strongly) incurved to the median part with in many specimens nearly parallel lateral margins; in apical third strongly tapering, hind margin distinctly medially excised over the whole length of the apical third, apical lobes laterally flattened, the apex round spatulate with a well-defined, slightly upwards-pointing, curved tooth at the lower end; in profile pointing backward. Styli (Fig. 174 ) short, thick, 1.1-1.6 × longer than wide, downwardly directed in lateral view, inserted quite far before apex of apical lobe.

Titillator (Figs 189 , 205 ) moderately sized; basal parts extending, strongly curved in the direction of the apical arms; fused part of apical arms broad at base not widening to the beginning of the unfused part of the apical arms; unfused part of apical arms hook-like, parallel or diverging and in lateral view in a 35-50 degrees angle curved upward to dorsum, wide at basis and evenly narrowing to tip; fused part of apical arms as long to longer than unfused part.

Colouration (in living specimens): general colouration in Andikithiran specimens dark brown (in several specimens chestnut brown) (Fig. 243 ), in Chania specimens green to light brown (Fig. 245 ). Head: frontal part below antennae and eyes in Andikithiran specimens creamy yellow-brownish with two larger inner and two smaller outer dark brown spots (Fig. 17 ) and often brownish speckled below the eyes, in Chania specimens bright green and likewise arranged and sized spots in black; border of frons with (lighter coloured) clypeus with dark transverse patches; upper part around eyes and antennal sockets black; behind both eyes and antennae two black bands separated from each other by a yellowish median line; occiput with black marking often provided with a thin lighter median line. Pronotum: dorsum dark brown-chestnut brown (Andikithira) to greenish or yellowish brown and often mottled (Chania) in first half with more (Andikithira) (Fig. 31 ) or less (Chania) extensive black marking; lateral lobes in upper part with black, ventrally not sharply delimited, longitudinal band, lower part pronotal lobes brownish to pinkish (Andikithira), green or yellow-white (Chania); in many specimens lower margin pronotal lobes in metazona yellowish. Elytra: visible parts not covered by the pronotum black or dark brown, covered part (lighter) brownish. Abdomen: first tergite dorsally black, other tergites completely dark brown to chestnut brown (Andikithira) or green to brownish often dorsally lighter coloured (Chania) and on both islands abdominal tergites sometimes mottled, anal tergite completely black; abdominal sternites pinkish brown (Andikithira) or yellowish brown (Chania). Cercus and subgenital plate: same (general) colour as body. Titillator: basal parts and unfused part of apical arms same colour as body, fused part of apical arms lighter coloured. Legs: same colour as body; fore and middle legs with many blackish to brownish stripes, spots, and markings; hind femur in the basal half dorsally with a longitudinal black to brownish stripe and also laterally on the outside in the middle part of its length; hind knees black.

Female. General appearance (Figs 230 , 231 ) as in male. Elytra completely covered by pronotum, only in some females scarcely protruding laterally.

Cercus short, conical with golden coloured short and long hairs, nearly straight, tapering apically; tip pointed, slightly bent inwards.

Subgenital plate (Figs 45 , 59 ) in ventral view generally wider than long; hind margin rounded, medially with a broadly rounded wide V-shaped excision half as long as the subgenital plate; basis concave with a shallow medial longitudinal ridge; in profile short triangular, apex rounded and not reaching or surpassing the proximal half of the gonangulum (Fig. 67 ).

Ovipositor nearly straight, only slightly upcurved near its apex, 1.5 to almost 2.0 × longer than pronotum.

Colouration generally as in male (Figs 244 , 246 ). Black marking of pronotum dorsally in prozona in most females less extensive as in males. First abdominal segment black; cercus, subgenital plate and ovipositor same colour as body (Andikithira) or yellowish brown with tip of ovipositor darker brown and laterally its medial part greyish brown.

Morphological variation found in E. francisae sp. nov. is elaborated in the Discussion.

Measurements.

See Tables 6 , 7 .

Bioacoustics.

Based upon the sound recordings of 15 specimens (153 syllables), the song of E. francisae sp. nov., 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. francisae sp. nov., the syllable duration is ~ 188 ms (Fig. 218 ). In the present recordings, the syllable repetition rate is slower than 0,5/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 francisae sp. nov. see Suppl. material 3.

Variation.

Within this new taxon, specimens from Andikithira are, as stated earlier, quite uniform in their morphological traits and colouration, while the populations on Crete incorporate more variation as the morphometric analyses in Tables 3 , 4 show. For the males this is especially the case in the presence/absence of tiny spines at the tip of the subgenital plate (compare Figs 216 , 217 ), the ratio length-width hind femur, the ratio length-width subgenital plate, the length of the incision of the subgenital plate. The females show most variation in the ratio length-width hind femur. Looking at the differences between the populations of Andikithira and those of western/southwestern Crete, the males and females of Andikithira in general have a larger body length and pronotum length; the males of Andikithira also possess a subgenital plate that is longer and wider than in those from Chania; the females of Andikithira have a subgenital plate that generally is wider than in those from Chania. Moreover, in females from Chania the length of the median incision of the hind margin of the subgenital plate is longer.

Differential diagnosis.

The new species differs from all the other species of the genus by the shape of the strikingly elongated male subgenital plate. Within the E. prasina group (male cerci of most taxa possess no tooth) the new species belongs to the E. latens subgroup as its preapically situated short styli are downward directed in lateral view. Eupholidoptera francisae sp. nov. seems most related to E. latens by the shape and proportions of the male subgenital plate with the apical lobes provided with a tooth at its tip, the proportions of the stylus, the shape of the titillator and the ratio height-length of the hind femur (see Tables 6 , 7 for measurements). A phylogeny based on molecular data also clearly separates E. francisae from E. latens (see discussion).

The male subgenital plate of E. francisae sp. nov. (Figs 146 , 160 ) is larger and more elongated than in E. latens (Figs 144 , 158 ). The stylus of E. francisae sp. nov. is 1.5 × longer than wide, while in E. latens it is 2-3 × longer than wide. The fused parts of the apical arms of the titillator of E. francisae sp. nov. (Fig. 189 ) are broad at base, not widening to the beginning of the unfused part, while in E. latens (Figs 186 , 187 ) they are narrow at base and clearly widening to the beginning of the unfused part. The unfused part of the apical arms of the titillator of E. francisae sp. nov. is not spine-like, straight and only slightly to moderately curved upward to the dorsum. In contrast, in E. latens the unfused part is spine-like and in most specimens strongly hooked upward to the dorsum.

The females of E. francisae sp. nov. differ from the other taxa in the genus by the shape and proportions of the subgenital plate (Figs 45 , 59 ). It can be distinguished from females of E. latens by the fact that in ventral view the incision of the hind margin is shaped in the form of a wide V, instead of slit-like or narrowly V-shaped as in E. latens ; in profile the apex of the female subgenital plate of E. francisae sp. nov. does not reach or surpass the proximal half of the gonangulum, while in E. latens the apex reaches the distal half of the gonangulum or even surpasses it. For more details differentiating E. francisae sp. nov. from other Cretan Eupholidoptera , see Table 5 .

Distribution.

This new taxon has been found on the island of Andikithira situated some 32 km NW of Crete and also in the western and southwestern part of Chania in western Crete (Fig. 254 ). Andikithira is a geographically isolated island halfway between the island of Kithira in the northwest and Crete in the southeast. It is a small, dry, and stony island (20.43 km2) rising to not more than 378 meters above sea level. The island has only few tens of residents and is hardly visited by tourists.

In Chania populations of E. francisae sp. nov. have been encountered west and southwest from the line of Gramvousa peninsula (northwest coast) to Livadas (near the south coast and situated 3-4 km west of the famous Samaria Gorge). Worth mentioning is also the fact that several males and females of E. francisae sp. nov. were caught in 1997 in a pitfall trap near Piso Moni Preveli. This location a long way to the east along the southern coast of the Rethimno region is ~ 60 km (in a straight line) east of Livadas. Piso Moni Preveli is also situated near the eastern (sic!) boundary of E. giuliae . Why E. francisae sp. nov. occurs here and has not been found on intermediate locations is puzzling. A revisit to this location to confirm its presence is necessary to rule out a mistake of mislabelling. For a complete list of localities, specimens and repositories see Suppl. material 1.

Habitat.

On Andikithira the species was found in phrygana and garrigue that cover a significant part of the entire island. Most specimens were collected as nymphs in Sarcopoterium spinosum that is present all over the island. The collecting sites are situated 50-150 m above sea level. But this species probably is present from sea level to the highest points of the island wherever phrygana and garrigue formations are present. On one of the collecting sites on Andikithira the new species was found together with the first specimen of Rhacocleis andikithirensis ( Tilmans et al. 2016). On both collecting sites on Andikithira also Poecilimon cretensis or a new taxon closely linked to it (pers. obs.) was present. In southwestern Chania E. francisae sp. nov. was not only found in low prickly shrublets in phrygana, but also frequently on tall shrubs of blackberry ( Rubus ).

Etymology.

Named in honour of Mrs. Francis Smid-Elbers, the late mother-in-law of the second author. Together with her husband Jacques Smid, she enthusiastically collected many interesting Orthoptera specimens in Greece, also from Crete. For instance, the paratype male and female of E. giuliae from 2.5 km E. of Argoules.

Phenology.

On Andikithira most specimens were collected as nymphs becoming adult in the period 22 May-10 June. In Chania collected nymphs became adult in the period 26 May-6 June and adults were collected in the period 23 May-21 June. Adults of Eupholidoptera francisae sp. nov. can thus be encountered from the end of May throughout June to July and possibly even later.

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 257. The Eupholidoptera giuliae - Eupholidoptera latens - Eupholidoptera francisae complex: geographic distribution of five morphological entities in the Eupholidoptera giuliae - Eupholidoptera latens - Eupholidoptera francisae complex: (1) eastern Chania + western Rethimno (E. giuliae), (2) northeastern Chania including Akrotiri peninsula (atypical E. giuliae), (3) northern and central Chania (E. latens), (4) western and southwestern Chania (E. francisae), (5) Andikithira (E. francisae).

Figure 258. Distribution results genetic analysis: geographic pattern of analysis of genetic data of Eupholidoptera populations in western Crete. yellow dots: E. francisae, purple dots: E. latens, blue dots: E. giuliae, light blue dots: atypical E. giuliae.

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

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 67, 68. Female subgenital plate in lateral view 67 Eupholidoptera francisae sp. nov. Anidhroi CT 2001.002.07 68 Eupholidoptera latens Prases DC-Ort 000512.

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 216, 217. Tip of male subgenital plate in ventral view 216 Eupholidoptera francisae sp. nov. paratype Anidhroi CT 2001.002.01 217 Eupholidoptera francisae sp. nov. paratype Anidhroi CT 2001.002.0.

Figures 218, 219. Oscillograms of Eupholidoptera 218 Single syllables of nine specimens of E. francisae sp. nov., timescale 500 ms and temperature 25 - 27.7 ° C a 2002.004.10 b 2002.004.09 c 2002.004.08 d 2002.004.07 e 2002.004.04 f 2002.004.07 g RMNH. 5087052 h 2001.002.02 i RMNH. 5106281 219 single syllables of one specimen of E. jacquelinae, timescale 500 ms and temperature 23.9 - 24.5 ° C a, b 2001.004.12.

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.

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.