Research Article |
Corresponding author: Luc Willemse ( luc.willemse@naturalis.nl ) Academic editor: Zhu-Qing He
© 2023 Luc Willemse, Jos Tilmans, Nefeli Kotitsa, Apostolos Trichas, Klaus-Gerhard Heller, Dragan Chobanov, Baudewijn Odé.
This is an open access article distributed under the terms of the Creative Commons Attribution License (CC BY 4.0), which permits unrestricted use, distribution, and reproduction in any medium, provided the original author and source are credited.
Citation:
Willemse L, Tilmans J, Kotitsa N, Trichas A, Heller K-G, Chobanov D, Odé B (2023) A review of Eupholidoptera (Orthoptera, Tettigoniidae) from Crete, Gavdos, Gavdopoula, and Andikithira. ZooKeys 1151: 67-158. https://doi.org/10.3897/zookeys.1151.97514
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Being nocturnal, hiding in prickly bushes and shrubs during the day, Eupholidoptera species in Crete and its neighbouring islands are easily overlooked, and until now our knowledge about their distribution was based on some thirty sightings across 11 species. In this paper results are presented of a study of Eupholidoptera specimens collected between 1987 and 2020 by hand-catches and pitfall and fermenting traps on the Greek islands of Crete, Gavdos, Gavdopoula, and Andikithira. Diagnostic features of all known species are presented and illustrated with stacked images. An updated key to all species is provided. Eupholidoptera francisae Tilmans & Odé, sp. nov. from Andikithira and southwestern Crete and Eupholidoptera marietheresae Willemse & Kotitsa, sp. nov. from Mt. Dikti are described. Female E. cretica, E. gemellata, and E. mariannae are described, and the female of E. astyla is redescribed. Bioacoustics for E. francisae Tilmans & Odé, sp. nov., E. giuliae, and E. jacquelinae are presented for the first time. Eupholidoptera smyrnensis is reported for the first time from Crete. A substantial amount of new distribution data for Eupholidoptera species on Crete is presented. The current distribution pattern and first analyses of phylogeny based on molecular data of Eupholidoptera species on Crete are discussed in relation to paleogeographical events.
Bioacoustics, faunistics, Greece, new species, phylogeography, systematics, traps
Eupholidoptera is a Mediterranean bush-cricket genus with 54 species belonging to the tribe Pholidopterini (
Species | Number of locations | Altitude (m) | Reference |
---|---|---|---|
E. annamariae | 2 | 50–200 |
|
E. astyla | 10 | 800–1850 |
|
E. cretica | 1 | not indicated |
|
E. feri | 1 | 1100 |
|
E. forcipata | 3 | 1700–1850 |
|
E. gemellata | 1 | 1650 |
|
E. giuliae | 3 | 0–175 |
|
E. jacquelinae | 1 | 50 |
|
E. latens | 7 | 500–1800 |
|
E. mariannae | 4(1) | 500–700 |
|
E. pallipes | 1 | 1600–1800 |
|
Hand catches included scanning larger shrubs (Rubus, Calicotome) in the morning to check for sunbathing individuals, walking through a terrain, and checking for individuals hiding in spiny bushes during the day. Additionally, singing males can be located using audio equipment late in the afternoon and in combination with a (head) light in the first half of the night. Specimens were killed using ethylacetate or potassium cyanide, eviscerated, the belly filled with cotton wool, and airdried.
Specimens found as nymphs were reared to adults. For this, cylindrical plastic containers with small twigs were used and nymphs were fed with oat flakes. In some cases, however, very young nymphs that, based on their dark colour pattern, were assumed to be Eupholidoptera turned out to belong to different genera (for instance Incertana Zeuner, 1941).
The traps were part of MSc and PhD studies aimed at the epigeal fauna of Crete and environmental monitoring programs, carried out by the Natural History Museum of the University of Crete (
Two types of traps were used, pitfall traps dug into the soil and fermenting traps placed higher up in the vegetation.
Pitfall traps (Fig.
Fermenting traps (Fig.
Specimens collected by hand were dried and pinned. Specimens caught by traps are stored in 70% alcohol except for a few which, after examination, were pinned. For the storage medium for each of the examined specimens see Suppl. material
Part of the DNA samples used for phylogenetic analysis derived from right mid legs from specimens collected in 2014, 2017, and 2019 (see Suppl. material
DNA was extracted from femoral muscles of Eupholidoptera specimens that were preserved either in alcohol (75% or 95%) or dry mounted. For recent and well-preserved specimens, a standard protocol for DNA isolation using ethanol precipitation was used. For dry-mounted and old, alcohol-preserved specimens from pitfall and fermenting traps DNA-isolation kits were used (Qiagen DNeasy Blood & Tissue Kits and Invitrogen purelink genomic DNA mini kit) following the manufacturers’ instructions.
Sequence data for one protein-coding mitochondrial gene (NADH dehydrogenase subunit 2 – NADH2) and one non-coding nuclear region (the internal transcribed spacers 1 and 2 together with the 5.8S rDNA gene in-between – ITS) where used. For amplification of NADH2 the primers used were TM-J210 AATTAAGCTAATGGGTTCATACCC (forward) and TW-N1284 AYAGCTTTGAARGYTATTAGTTT (reverse) (
Polymerase chain reactions were performed using Thermo Scientific DreamTaq Hot Start Master Mix according to the manufacturer’s instructions. Temperature cycling for the NADH2 followed
Additional sequences for both loci were obtained from GenBank (https://www.ncbi.nlm.nih.gov/genbank/).
Obtained sequences were trimmed, assembled, and visually checked using CodonCode Aligner v. 8.0.2 (CodonCode, Dedham, MA, USA). All protein-coding sequences were checked for numt possibility and unique haplotypes were selected using DAMBE 7.2.152 (Xia, 2018). Sequence alignments were performed in MEGA-X (
Bayesian inference (BI) phylogenetic analysis was performed on the concatenated dataset (NADH2 + ITS) using Mr. Bayes v. 3.2.7 (
Coordinates are presented in decimal degrees (DD) and may therefore differ from collecting labels that use degrees, minutes and seconds or degrees and minutes (DMS). For locations for which no coordinates were available Google Earth has been used in which case coordinates have been placed between square brackets “ []”.
Titillators, once removed, were cleared in a KOH solution and fixed with glue on a small board, pinned under the specimen or in case of E. latens from Rhodopos, transferred to glycerol (
For song recordings several digital recorder systems (digital tape, solid state memory and computer hard disk) and microphones were used. Usually a frequency response better than 50–20,000 Hz were achieved. Most recordings were made indoors in a lab or room, in the evening or night, using (partly) open containers, frequently housed in an anechoic cupboard, with the microphone at 3–15 cm distance. The air temperature during recording in room or studio was between 15 °C and 27 °C. We did not attempt to correct for the possible body temperature of the animals, which may well have been below or above the measured room temperature. Specific data for the individual recordings can be found in Suppl. material
Song analysis of the digital recordings has been performed using Wavelab software (www.steinberg.net). Oscillograms have been prepared using Praat software (www.praat.org).
Bioacoustic terminology: calling song – the song produced by an isolated male; syllable – the sound produced by one opening-and-closing movement of the tegmina; hemisyllable – the sound produced by the opening or closing movement of the tegmina; syllable period – period from one syllable beginning to the next. Syllable repetition rate – the number of syllables produced per second.
For stacked images, a Zeiss SteREO Discovery V20 stereomicroscope was used, combined with a Zeiss AxioCam MRc5 microscope camera. The habitus photographs were taken with a NIKON D5600 with a sigma 105 mm macrolens and a Canon EOS 5D digital camera using a Canon zoom lens EF 28–90 mm F 4–5.6 with three combined Hama Close-Up lenses 1, 2 + 4×.
Figs
A list summarising all known localities, specimens, and repositories is presented in Suppl. material
BMNH British Museum of Natural History, London, UK;
CH collection Klaus-Gerhard Heller, Triesdorf, Germany;
CMUP collection Bruno Massa, University of Palermo, Palermo, Italy;
CT collection Jos Tilmans, Wassenberg-Rothenbach, Germany;
MfNB Museum für Naturkunde, Berlin, Germany;
obs. observation (specimen not collected);
Table
Published and unpublished records (period 1987–2020) of Eupholidoptera from Crete and adjoining islands.
Altitudinal range (m) | Published locations (Table |
Tilmans 1987–2019 | Trapping program 1987–2019 |
|
Observations 2012–2020 | Heller 2016 | Willemse and Zacharopoulou 2017 | Chobanov et al. 2018 | Tilmans and Willemse 2019 | Total | |
---|---|---|---|---|---|---|---|---|---|---|---|
E. annamariae | 5–550 | 2 | 5 | 3 | 0 | 0 | 0 | 2 | 2 | 0 | 14 |
E. astyla | 5–1850 | 10 | 6 | 24 | 0 | 0 | 1 | 3 | 1 | 0 | 45 |
E. cretica | 1165–1234 | 1 | 0 | 4 | 0 | 0 | 0 | 0 | 0 | 0 | 5 |
E. feri | 1100 | 1 | 0 | 0 | 0 | 0 | 0 | 0 | 0 | 0 | 1 |
E. forcipata | 1350–2225 | 3 | 2 | 3 | 0 | 0 | 0 | 0 | 0 | 0 | 8 |
E. francisae sp. nov. | 1–835 | 0 | 2 | 1 | 0 | 0 | 0 | 2 | 2 | 9 | 16 |
E. gemellata | 1650–1910 | 1 | 0 | 1 | 0 | 0 | 0 | 0 | 0 | 0 | 2 |
E. giuliae | 0–525 | 3 | 4 | 9 | 0 | 7 | 1 | 3 | 1 | 4 | 32 |
E. jacquelinae | 30–270 | 1 | 0 | 8 | 0 | 1 | 0 | 0 | 0 | 0 | 10 |
E. latens | 20–1815 | 7 | 3 | 4 | 1 | 0 | 1 | 1 | 2 | 2 | 21 |
E. mariannae | 0–1475 | 4 | 2 | 9 | 0 | 0 | 0 | 2 | 0 | 0 | 17 |
E. marietheresae sp. nov. | 1715 | 0 | 0 | 1 | 0 | 0 | 0 | 0 | 0 | 0 | 1 |
E. pallipes | 1600–2440 | 1 | 0 | 1 | 0 | 1 | 0 | 0 | 0 | 0 | 3 |
E. smyrnensis | 25–340 | 0 | 0 | 1 | 0 | 0 | 0 | 1 | 0 | 1 | 3 |
Total | 34 | 23 | 69 | 1 | 9 | 3 | 14 | 8 | 16 | 178 |
1 | Frons with tiger-pattern (Fig. |
smyrnensis |
– | Frons with isolated dark spots or black crossband (Figs |
2 |
2 | Females | 3 |
– | Males | 15 |
3 | Subgenital plate with pair of elongate concavities divided by median ridge, apical lobes touching (Fig. |
annamariae |
– | Subgenital plate convex or proximally concave or with pits, apical lobes separated (Figs |
4 |
4 | Gavdos or Gavdopoula | jacquelinae |
– | Crete or Andikithira | 5 |
5 | Subgenital plate wider to much wider than long (Figs |
6 |
– | Subgenital plate as wide as long or elongated (Figs |
10 |
6 | Hind margin of subgenital plate with wide and deep median excision (Fig. |
cretica |
– | Hind margin of subgenital plate differently formed (Figs |
7 |
7 | Hind margin of subgenital plate medially concave, without excision (Fig. |
gemellata |
– | Hind margin of subgenital plate with distinct excision (Figs |
8 |
8 | Subgenital plate with apical lobes rounded (Fig. |
francisae sp. nov. |
– | Subgenital plate with apical lobes pointed (Figs |
9 |
9 | Subgenital plate 2.0× wider than long, proximal pits small, widely separated (Fig. |
forcipata |
– | Subgenital plate 1.4× wider than long, proximal pits large, closer to each other (Fig. |
marietheresae sp. nov. |
10 | Front of head with large black patches (Fig. |
pallipes |
– | Front of head with black dots (Figs |
11 |
11 | Apex subgenital plate in profile (measured in a straight line parallel to the ovipositor) not reaching or surpassing proximal half of gonangulum (Fig. |
francisae sp. nov. |
– | Apex subgenital plate in profile (measured in a straight line parallel to the ovipositor) reaching or surpassing distal half of gonangulum (Fig. |
12 |
12 | Subgenital plate proximally convex, flattened or slightly concave (Fig. |
astyla |
– | Subgenital plate proximally with one wide or two separate pits (Figs |
13 |
13 | Apical lobes of subgenital plate not well produced, rectangular (Fig. |
latens |
– | Apical lobes of subgenital plate more produced, rectangular to acute (Figs |
14 |
14 | Subgenital plate proximally with two distinct pits separated by a median keel (Fig. |
giuliae |
– | Subgenital plate proximally with a single or two pits (Figs |
mariannae |
– | (Mt. Dikti, Katharo plains) | feri |
15 | Styli pointing backwards (Figs |
16 |
– | Styli pointing downward or inward (Figs |
21 |
16 | Cercus with subbasal side tooth (Figs |
17 |
– | Cercus unarmed (Figs |
18 |
17 | Pronotum pale (Fig. |
pallipes |
– | Pronotum with black band or patches (Fig. |
gemellata |
18 | Styli very long, 5–6× longer than wide (Figs |
19 |
– | Styli short, 2× longer than wide (Figs |
20 |
19 | Subgenital plate with long spine at base of stylus (Fig. |
jacquelinae |
– | Subgenital plate without a spine at base of stylus (Fig. |
cretica |
20 | Hind margin of anal tergite from the cercus downward straight, side flaps gradually narrowing (Fig. |
forcipata |
– | Hind margin of anal tergite from the cercus downward S-curved, side flaps first widening before narrowing (Fig. |
marietheresae sp. nov. |
21 | Anal tergite distally extended into two very long, spined hooks pointing downward (Fig. |
annamariae |
– | Anal tergite distally extended into short pointed lobes (Figs |
22 |
22 | Pointed lobes of anal tergite close together (Figs |
23 |
– | Pointed lobes of anal tergite widely separated (Figs |
25 |
23 | Cercus unarmed (Fig. |
astyla |
– | Cercus with subbasal side tooth (Figs |
24 |
24 | Cercus conical (Fig. |
feri |
– | Cercus flattened and widened proximally (Fig. |
mariannae |
25 | Subgenital plate as wide as long (Fig. |
giuliae |
– | Subgenital plate elongate, tapering toward the apex (Figs |
26 |
26 | Tips apical lobes subgenital plate with a tooth (Figs |
francisae sp. nov. |
– | Tips apical lobes subgenital plate always without a tooth (Figs |
latens |
The taxonomic treatment contains short diagnostics, illustrated with stacked images, for all species until now reported from Crete and its adjoining islands. New taxa as well as previously unknown sexes are described in more detail.
Eupholidoptera annamariae
Morphological description.
Bioacoustics.
1 ♂, 1 ♀ (paratypes); 23 ♂, 24 ♀ (for details see Suppl. material
Frontal part of head (Fig.
Colour pattern of head, frontal view 11 Eupholidoptera smyrnensis ♀ Makrigiannis RMNH5014918 12 Eupholidoptera gemellata ♂ Mt. Idhi
Colour pattern of pronotum, dorsal view 25 Eupholidoptera smyrnensis ♂ Makrigiannis
Female subgenital plate in ventral view 39 Eupholidoptera smyrnensis Makrigiannis
Female subgenital plate in lateral view 53 Eupholidoptera smyrnensis Makrigiannis
See Tables
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 (
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.
Males differ from congenerics in the anal tergite (Figs
Male anal tergite in dorsal view 69 Eupholidoptera smyrnensis Makrigiannis
Male anal tergite in caudal view 83 Eupholidoptera smyrnensis Makrigiannis
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
Male cercus in dorsal view 111 Eupholidoptera smyrnensis Makrigiannis RMNH5087053 112 Eupholidoptera gemellata Mt. Idhi FC1651
Male cercus in lateral view 126 Eupholidoptera smyrnensis Makrigiannis
Male subgenital plate in ventral view 140 Eupholidoptera smyrnensis Lagkada
Male subgenital plate in lateral view 154 Eupholidoptera smyrnensis Lagkada
Male stylus in lateral view 168 Eupholidoptera smyrnensis Makrigiannis
Male titillator in ventral view 182 Eupholidoptera smyrnensis Makrigiannis RMNH5087053 183 Eupholidoptera gemellata Mt. Idhi Amariou FC1651
Male titillator in lateral view 198 Eupholidoptera smyrnensis Makrigiannis RMNH5087053 199 Eupholidoptera gemellata Mt. Idhi Amariou FC1651
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 (
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.
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.
Pholidoptera astyla
Eupholidoptera astyla
(Ramme, 1927);
Morphological description.
Bioacoustics.
1 ♂ (paratype); 81 ♂, 59 ♀ (for details see Suppl. material
Frontal part of head (Fig.
In 1927 E. astyla was described based on a single male from Naxos and three females and the male abdomen from Crete (
Female.
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),
General appearance (Figs
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].
See Tables
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 (
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.
Males differ from congenerics in the strongly asymmetrical, thickened and wrinkled apical arms of the titillator (Figs
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.
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.
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.
Eupholidoptera cretica Ramme, 1951: 202.
Morphological description.
Eupholidoptera cretica was described after a single male, collected 13 June 1942 by K. Zimmermann. This, most likely, is the mammologist who worked on the mammals of Crete and published a review of his observations including a map (
1 ♂, 2 ♀ (for details see Suppl. material
Frontal part of head (Fig.
Female. Examined specimens. 2♀: CHANIA: Lefka Mt., above Omalos –
General appearance (Figs
See Tables
The song of this species has not yet been recorded.
Males differ from congenerics in the stout, unarmed, inward curved cercus (Figs
Besides the type location which is not exactly traceable, only known from two spots on Mt. Lefka, one in northwest near the Omalos plateau and Samaria gorge and a second along the southeastern slopes above the villages of Anopoli and Limnia (Fig.
The area around the Omalos plateau where the species was trapped is described as Cupressus forest. The species has been trapped in fermenting traps placed above the ground in shrubs, indicating E. cretica like E. smyrnensis, E. mariannae as well as E. jacquelinae but contrary to most other Cretan species, actually lives in such shrubs and not in small prickly bushes on the ground.
Still very little is known. The first male was caught on 13 June 1942. Specimens being caught in traps were found in traps operative between 31 July and 19 October. The recorded altitudes where the species was found are between 1165 m and 1235 m.
Eupholidoptera rammei Willemse & Heller, 2001: 333.
Eupholidoptera feri Koçak & Kemal, 2010: 7.
Morphological description.
Bioacoustics.
Holotype, allotype (for details see Suppl. material
Frontal part of head (Fig.
See Tables
Based upon the sound recordings of 1 specimen (10 syllables measured), the song of E. feri, 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. The syllable duration is ~ 271 ms, recorded at 15 °C, with a syllable rate up to ~ 1/s. Published records (
Males differ from congenerics in the stout, cylindrical cerci (Figs
Only known from the Katharo plain in the eastern offshoots of Mt. Dikti, in the western part of the Lasithi district (Fig.
The type specimens were collected 1–2 m high in a Quercus shrub in the Katharo plain, part of which is used for cultivation (vineyards), the rest consists of bare grounds.
The Katharo plain lies at an altitude of 1100 m. The type specimens were collected in late August.
Eupholidoptera forcipata Willemse & Kruseman, 1976: 131.
Morphological description.
Bioacoustics.
Holotype, allotype, 8 ♂, 9 ♀ (paratypes); 12 ♂, 5 ♀ (for details see Suppl. material
Frontal part of head (Fig.
See Tables
Based upon the sound recordings of one specimen (30 syllables measured), the song of E. forcipata, 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. forcipata, the syllable duration is ~ 525 ms, with a syllable rate up to somewhat less than 1/s. The syllable duration easily discerns the song of this species from the other known songs of Eupholidoptera from Crete. Published records (
Males differ from congenerics in the pointed backward and downward extended widened apical lobes of the anal tergite (Figs
Only known from higher altitudes on Mt. Psiloritis, central Crete (Fig.
The species lives at high altitudes in subalpine phrygana in low prickly bushes (e.g., Astragalus) in which it hides during the day.
The species occurs between 1350 m and 2225 m. Adults have been collected by hand at the end of July and during the first half of August. Trap catches indicate they are still active up to September and possibly October.
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.
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 ♀ (
Male. General appearance (Figs
Pronotum (Fig.
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
Cerci (Figs
Subgenital plate (Figs
Titillator (Figs
Colouration (in living specimens): general colouration in Andikithiran specimens dark brown (in several specimens chestnut brown) (Fig.
Female. General appearance (Figs
Cercus short, conical with golden coloured short and long hairs, nearly straight, tapering apically; tip pointed, slightly bent inwards.
Subgenital plate (Figs
Ovipositor nearly straight, only slightly upcurved near its apex, 1.5 to almost 2.0× longer than pronotum.
Colouration generally as in male (Figs
Morphological variation found in E. francisae sp. nov. is elaborated in the Discussion.
See Tables
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.
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
Males | Euph. latens | Euph. francisae | Euph. francisae only Andikithira | Euph. francisae only Chania |
---|---|---|---|---|
length body | n = 9 | n = 38 | n = 11 | n = 27 |
min. – max. | 18.9–27.1 | 19.0–28.8 | 20.9–28.3 | 19.0–28.8 |
mean ± SD | 22.4 ± 2.58 | 24.1 ± 2.66 | 25.6 ± 2.58 | 23.5 ± 2.50 |
length pronotum | n = 9 | n = 38 | n = 11 | n = 27 |
min. – max. | 8.4–9.9 | 8.0–10.7 | 9.0–10.7 | 8.0–10.4 |
mean ± SD | 9.1 ± 0.53 | 9.5 ± 0.65 | 10.0 ± 0.53 | 9.3 ± 0.59 |
length hind femur | n = 13 | n = 37 | n = 10 | n = 27 |
min. – max. | 17.0–22.7 | 19.0–23.1 | 19.0–22.8 | 19.7–23.1 |
mean ± SD | 19.3 ± 2.28 | 21.1 ± 0.93 | 20.7 ± 1.10 | 21.2 ± 0.83 |
width hind femur | n = 13 | n = 37 | n = 10 | n = 27 |
min. – max. | 3.9–4.8 | 3.7–4.9 | 4.0–4.6 | 3.7–4.9 |
mean ± SD | 4.3 ± 0.23 | 4.3 ± 0.25 | 4.4 ± 0.18 | 4.3± 0.27 |
ratio length-width hind femur | n = 13 | n = 37 | n = 10 | n = 27 |
min. – max. | 4.14–5.28 | 4.47–5.49 | 4.52–5.07 | 4.47–5.49 |
mean ± SD | 4.53 ± 0.37 | 4.88 ± 0.25 | 4.74 ± 0.16 | 4.93 ± 0.26 |
length subg. plate | n = 12 | n = 34 | n = 11 | n = 23 |
min. – max. | 3.75–6.30 | 4.25–5.90 | 5.35–5.90 | 4.25–5.80 |
mean ± SD | 4.45 ± 0.74 | 5.08 ± 0.49 | 5.57 ± 0.17 | 4.85 ± 0.41 |
width subg. plate | n = 12 | n = 34 | n = 11 | n = 23 |
min. – max. | 2.20–5.00 | 2.00–3.85 | 2.95–3.85 | 2.00–3.75 |
mean ± SD | 3.21 ± 0.78 | 3.13 ± 0.48 | 3.43 ± 0.32 | 2.99 ± 0.48 |
ratio length-width subg. plate | n = 12 | n = 34 | n = 11 | n = 23 |
min. – max. | 0.84–2.05 | 1.13–2.44 | 1.48–1.90 | 1.13–2.44 |
mean ± SD | 1.44 ± 0.32 | 1.66 ± 0.30 | 1.63 ± 0.15 | 1.67 ± 0.35 |
length incision subg. plate | n = 12 | n = 32 | n = 11 | n = 21 |
min. – max. | 1.15–1.60 | 1.20–1.90 | 1.50–1.85 | 1.20–1.90 |
mean ± SD | 1.33 ± 0.15 | 1.53 ± 0.21 | 1.67 ± 0.13 | 1.46 ± 0.21 |
Measurements (in mm) and biometrics of female E. latens and E. francisae.
Females | Euph. latens | Euph. francisae | Euph. francisae only Andikithira | Euph. francisae only Chania |
length body | n = 8 | n = 38 | n = 11 | n = 27 |
min. – max. | 17.5–23.3 | 19.5–31.5 | 23.7–31.5 | 19.5–26.9 |
mean ± SD | 20.3 ± 2.32 | 24.1 ± 2.43 | 26.3 ± 1.93 | 23.2 ±1.98 |
length pronotum | n = 8 | n = 38 | n = 11 | n = 27 |
min. – max. | 8.1–9.4 | 8.5–10.5 | 8.5–10.5 | 8.6–10.4 |
mean ± SD | 8.7 ± 0.46 | 9.6 ± 0.56 | 9.8 ± 0.65 | 9.5 ± 0.49 |
length ovipositor | n = 8 | n = 38 | n = 11 | n = 27 |
min. – max. | 13.8–16.4 | 13.8–19.3 | 15.8–18.7 | 13.8–19.3 |
mean ± SD | 14.9 ± 0.76 | 16.6 ± 1.38 | 17.5 ± 0.89 | 16.2 ± 1.39 |
ratio length ovip. pronot. | n = 8 | n = 38 | n = 11 | n = 27 |
min. – max. | 1.64–1.79 | 1.52–1.99 | 1.61–1.99 | 1.52–1.91 |
mean ± SD | 1.72 ± 0.05 | 1.73 ± 0.11 | 1.79 ± 0.12 | 1.71 ± 0.11 |
length hind femur | n = 9 | n = 37 | n = 11 | n = 26 |
min. – max. | 17.1–22.9 | 20.9–24.9 | 20.0–21.7 | 20.9–24.9 |
mean ± SD | 18.8 ± 1.93 | 22.2 ± 1.04 | 21.2 ± 0.61 | 22.6 ± 0.90 |
width hind femur | n = 9 | n = 37 | n = 11 | n = 26 |
min. – max. | 3.9–4.8 | 4.1–5.1 | 4.1–4.7 | 4.1–5.1 |
mean ± SD | 4.3 ± 0.27 | 4.5 ± 0.24 | 4.5 ± 0.20 | 4.5 ± 0.25 |
ratio length-width hind femur | n = 9 | n = 37 | n = 11 | n = 26 |
min. – max. | 3.98–4.77 | 4.55–5.62 | 4.55–5.02 | 4.60–5.62 |
mean ± SD | 4.40 ± 0.25 | 4.93 ± 0.28 | 4.76 ± 0.17 | 5.00 ± 0.28 |
length subg. plate | n = 9 | n = 35 | n = 11 | n = 24 |
min. – max. | 2.10–3.75 | 1.75–2.70 | 1.90–2.60 | 1.75–2.70 |
mean ± SD | 2.68 ± 0.51 | 2.19 ± 0.23 | 2.18 ± 0.25 | 2.19 ± 0.23 |
width subg. plate | n = 9 | n = 35 | n = 11 | n = 24 |
min. – max. | 2.10–3.20 | 1.95–3.40 | 2.20–3.40 | 1.95–2.90 |
mean ± SD | 2.62 ± 0.39 | 2.53 ± 0.41 | 2.93 ± 0.39 | 2.34 ± 0.27 |
ratio length-width subg. plate | n = 9 | n = 35 | n = 11 | n = 24 |
min. – max. | 0.86–1.44 | 0.59–1.23 | 0.59–1.00 | 0.75–1.23 |
mean ± SD | 1.03 ± 0.17 | 0.89 ± 0.15 | 0.76 ± 0.14 | 0.95 ± 0.12 |
length incision subg. plate | n = 9 | n = 35 | n = 11 | n = 24 |
min. – max. | 1.15–1.70 | 0.90–1.60 | 0.90–1.25 | 0.95–1.60 |
mean ± SD | 1.32 ± 0.18 | 1.16 ± 0.16 | 1.08 ± 0.12 | 1.20 ± 0.17 |
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
The male subgenital plate of E. francisae sp. nov. (Figs
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
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
Diagnostic characters and character-states for Eupholidoptera species from Crete, Andikithira, Gavdos, and Gavdopoula.
Structure | 1 | 2 | 3 | 4 | 5 | 6 | 7 | |||||||||||||||
---|---|---|---|---|---|---|---|---|---|---|---|---|---|---|---|---|---|---|---|---|---|---|
Character | a | b | c | a | b | c | a | b | a | b | c | d | a | b | a | b | c | d | a | b | c | d |
Species | ||||||||||||||||||||||
annamariae | 1 | 3 | 1 | 2 | 2 | 2 | 2 | 1 | 1 | 2 | 1 | 1 | 1 | 1 | 2 | 1 | 1 | 1 | 1–3 | 2 | 1 | 1 |
astyla | 1 | 3 | 1 | 1 | 1 | 1 | 3 | 1 | 1 | 1 | 1 | 1 | 1 | 1 | 3 | 1 | 3 | 1 | 3–4 | 2 | 1 | 1 |
cretica | 1 | 1 | 1 | 1 | 3 | 3 | 2 | 1 | 1 | 2 | 1 | 1 | 3 | 2 | 1 | 1 | 2 | 2 | 1 | 3 | 2 | 1 |
feri | 1 | 3 | 1 | 1 | 1 | 1 | 2 | 3 | 1 | 1 | 1 | 1 | 1 | 1 | 2 | 1 | 2 | 1 | 4 | 2 | 1 | 2 |
forcipata | 1 | 1 | 1 | 2 | 3 | 1 | 1 | 1 | 1–2 | 3 | 1 | 1 | 2 | 2 | 1 | 1 | 3 | 1 | 1 | 1 | 2 | 3 |
francisae | 1 | 2 | 1 | 1 | 2 | 1 | 3 | 1 | 3 | 2 | 1 | 1 | 1 | 1 | 1 | 1 | 3 | 2 | 1–3 | 3 | 2 | 2–3 |
gemellata | 3 | 3 | 3 | 1 | 1 | 2 | 2 | 3 | 1–3 | 2 | 2 | 2 | 2 | 2 | 1 | 2 | 2 | 2 | 1 | 1 | 2 | 1 |
giuliae | 1 | 2 | 1 | 1 | 2 | 1 | 3 | 1 | 1–3 | 1 | 1 | 1 | 1 | 3 | 1 | 1 | 3 | 2 | 2–4 | 3 | 1 | 3 |
jacquelinae | 1 | 1 | 1 | 2 | 2 | 1 | 3 | 1 | 3 | 3 | 2 | 1 | 3 | 2 | 1 | 1 | 3 | 2 | 3–4 | 3 | 1 | 1 |
latens | 1 | 2 | 1 | 1 | 2 | 1 | 3 | 1 | (1)-3 | 2 | 1 | 1 | 2 | 1 | 1 | 1 | 3 | 2 | 2 | 3 | 1 | 2–3 |
mariannae | 1 | 3 | 1 | 1 | 1 | 3 | 1 | 3 | 2 | 1 | 1 | 1 | 1 | 1 | 2 | 1 | 3 | 1 | 3–4 | 2 | 1 | 2–3 |
marietheresae | 3 | 1 | 1 | 2 | 3 | 1 | 1 | 1 | 1 | 3 | 1 | 1 | 2 | 2 | 1 | 1 | 3 | 1 | 1 | 2 | 2 | 3 |
pallipes | 3 | 3 | 3 | 1 | 1 | 2 | 2 | 3 | 2 | 2 | 2 | 2 | 2 | 2 | 1 | 2 | 1 | 2 | 2 | 1 | 2 | 1 |
smyrnensis | 2 | 1 | 2 | 1 | 3 | 1 | 1 | 2 | 2 | 3 | 3 | 2 | 2 | 2 | 1 | 1 | 3 | 2 | 3 | 2 | 2 | 1 |
1. Male and Female colouration | ||||||||||||||||||||||
a. frons | 1. dots (Fig. |
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b. pronotal disc | 1. pale (Fig. |
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c. abdomen | 1. pale (Fig. |
|||||||||||||||||||||
2. Male anal tergite | ||||||||||||||||||||||
a. extended backwards | 1. not/hardly; 2. distinctly | |||||||||||||||||||||
b. excision tips | 1. narrow; 2. intermediate; 3. wide | |||||||||||||||||||||
c. direction tips | 1. downward; 2. inward; 3. forward | |||||||||||||||||||||
3. Male cercus | ||||||||||||||||||||||
a. length-width ratio | 1. < 5; 2. 5–6.5; 3. > 6.5 | |||||||||||||||||||||
b. side tooth | 1. missing; 2. basal; 3. sub-basal | |||||||||||||||||||||
4. Male subgenital plate | ||||||||||||||||||||||
a. length-width ratio | 1. < 1; 2. ca. 1; 3. > 1 | |||||||||||||||||||||
b. length excision-total length ratio | 1. 0.1–0.25; 2. 0.3–0.4; 3. 0.5–0.6 | |||||||||||||||||||||
c. spine | 1. absent; 2. one; 3. two | |||||||||||||||||||||
d. protuberance | 1. absent; 2. present | |||||||||||||||||||||
5. Male styli | ||||||||||||||||||||||
a. length-width ratio | 1. 1.0–2.0; 2.0–3.0; 3. > 4.0 | |||||||||||||||||||||
b. direction | 1. downward; 2. backward; 3. inward | |||||||||||||||||||||
6. Male titillator | ||||||||||||||||||||||
a. symmetry | 1. symmetrical; 2. subsymmetrical; 3. asymmetrical | |||||||||||||||||||||
b. basal arms | 1. long; 2. short | |||||||||||||||||||||
c. apical arms | 1. merged; 2. largely merged; 3. free for > ⅓ | |||||||||||||||||||||
d. apical arms | 1. basal half stalk-like; 2. basal half wide | |||||||||||||||||||||
7. Female subgenital plate | ||||||||||||||||||||||
a. length-width ratio | 1. < 0.75; 2. 0.75–0.90; 3. 0.90–1.10; 4. >1.10 | |||||||||||||||||||||
b. length excision-total length ratio | 1. < 0.25; 2. 0.25–0.33; 3. > 0.33 | |||||||||||||||||||||
c. medial excision | 1. narrow; 2. wide | |||||||||||||||||||||
d. proximally | 1. convex; 2. concave; 3. with 2 concavities |
Males | Body length | Pronotum length | Hind femur length | Hind femur width | Ratio length-width hind femur | Number of teeth |
---|---|---|---|---|---|---|
annamariae | n = 23 | n = 23 | n = 23 | n = 23 | n = 23 | n = 1 |
mean ± SD | 28.3±2.1 | 11.3±0.5 | 21.3±0.8 | 4.8±0.2 | 4.43±0.16 | |
min. – max. | 25.0–32.2 | 10.5–13.1 | 19.6–22.8 | 4.4–5.1 | 4.16–4.81 | 109 |
astyla | n = 79 | n = 79 | n = 70 | n = 70 | n = 70 | n = 2 |
mean ± SD | 25.5±2.4 | 9.9±1.0 | 19.5±2.0 | 4.5±0.4 | 4.34±0.17 | |
min. – max. | 21.0–30.8 | 8.2–12.2 | 16.2–23.0 | 3.9–5.3 | 3.77–4.74 | 101–105 |
cretica | n = 1 | n = 1 | n = 1 | n = 1 | n = 1 | n = 1 |
mean ± SD | n.a. | n.a. | n.a. | n.a. | n.a. | |
min. – max. | 21.7 | 8.8 | 22.1 | 4.4 | 5.04 | 107 |
feri | n = 1 | n = 1 | n = 1 | n = 1 | n = 1 | n = 1 |
mean ± SD | n.a. | n.a. | n.a. | n.a. | n.a. | |
min. – max. | 26.8 | 8.8 | 18.6 | 4.0 | 4.61 | 100 |
forcipata | n = 20 | n = 20 | n = 20 | n = 20 | n = 20 | n = 1 |
mean ± SD | 23.4±1.6 | 9.5±0.5 | 17.8±0.7 | 4.3±0.2 | 4.18±0.11 | |
min. – max. | 20.8–26.7 | 8.6–10.5 | 16.6–19.6 | 4.0–4.7 | 3.96–4.43 | 193 |
francisae | n = 38 | n = 38 | n = 37 | n = 37 | n = 37 | n = 19 |
mean ± SD | 24.1±2.7 | 9.5±0.7 | 21.1±0.9 | 4.3±0.3 | 4.88±0.25 | 119±10.3 |
min. – max. | 19.0–28.8 | 8.0–10.7 | 19.0–23.1 | 3.7–4.9 | 4.47–5.49 | 96–138 |
gemellata | n = 3 | n = 4 | n = 4 | n = 4 | n = 4 | n = 1 |
mean ± SD | 21.1±3.9 | 7.4±0.3 | 17.1±0.2 | 4.0±0.1 | 4.32±0.10 | |
min. – max. | 16.8–24.2 | 7.0–7.7 | 16.8–17.3 | 3.9–4.1 | 4.17–4.45 | 101 |
giuliae | n = 52 | n = 52 | n = 52 | n = 52 | n = 52 | n = 1 |
mean ± SD | 24.5±3.2 | 10.2±0.5 | 21.7±0.8 | 4.7±0.2 | 4.66±0.24 | |
min. – max. | 17.2–30.2 | 9.1–11.4 | 20.0–23.8 | 4.0–5.0 | 4.26–5.43 | 106 |
jacquelinae | n = 6 | n = 7 | n = 7 | n = 7 | n = 7 | n = 1 |
mean ± SD | 27.4±2.5 | 10.6±0.3 | 24.8±1.1 | 4.8±0.2 | 5.14±0.36 | |
min. – max. | 25.3–32.1 | 10.2–11.2 | 23.8–26.4 | 4.4–5.1 | 4.86–5.98 | 144 |
latens | n = 9 | n = 9 | n = 13 | n = 13 | n = 13 | n = 4 |
mean ± SD | 22.4±2.6 | 9.1±0.5 | 19.3±2.3 | 4.3±0.2 | 4.53±0.37 | 113±7.1 |
min. – max. | 18.9–27,1 | 8.4–9.9 | 17.0–22.7 | 3.9–4.8 | 4.14–5.28 | 108–123 |
mariannae | n = 9 | n = 9 | n = 9 | n = 9 | n = 9 | n = 4 |
mean ± SD | 24.4±2.2 | 10.1±0.4 | 21.2±0.9 | 4.6±0.3 | 4.65±0.17 | |
min. – max. | 21.5–28.9 | 9.5–10.9 | 19.8–22.3 | 4.2–4.9 | 4.36–5.00 | 89–105 |
marietheresae | n = 5 | n = 5 | n = 5 | n = 5 | n = 5 | n = 2 |
mean ± SD | 24.9±2.7 | 9.4±0.6 | 17.1±0.6 | 4.0±0.2 | 4.30±0.14 | |
min. – max. | 21.4–28.5 | 8.9–10.3 | 16.4–18.1 | 3.8–4.2 | 4.04–4.45 | 211–213 |
pallipes | n = 6 | n = 6 | n = 6 | n = 6 | n = 6 | n = 1 |
mean ± SD | 20.3±1.0 | 7.7±0.3 | 16.9±0.6 | 4.1±0.1 | 4.18±0.13 | |
min. – max. | 19.3–21.9 | 7.4–8.1 | 16.5–18.0 | 3.9–4.2 | 3.92–4.32 | 94 |
smyrnensis | n = 7 | n = 7 | n = 7 | n = 7 | n = 7 | n = 1 |
mean ± SD | 20.2±1.6 | 9.8±0.4 | 23.3±0.6 | 4.5±0.1 | 5.24±0.19 | |
min. – max. | 17.5–22.6 | 9.5–10.3 | 22.4–24.0 | 4.3–4.6 | 4.92–5.48 | 100 |
Females | Body length | Pronotum length | Hind femur length | Hind femur width | Ratio length-width hind femur | Ovipositor length | Ratio length ovipositor-length pronotum |
---|---|---|---|---|---|---|---|
annamariae | n = 24 | n = 24 | n = 24 | n = 24 | n = 24 | n = 24 | n = 24 |
mean ± SD | 25.6±2.6 | 10.8±0.5 | 21.7±0.6 | 4.9±0.2 | 4.48±0.14 | 19.0±1.1 | 1.76±0.09 |
min. – max. | 21.7–32.2 | 10.2–12.1 | 20.4–22.8 | 4.6–5.2 | 4.20–4.81 | 17.0–21.2 | 1.63–2.05 |
astyla | n = 59 | n = 59 | n = 59 | n = 59 | n = 59 | n = 59 | n = 59 |
mean ± SD | 23.8±2.8 | 9.6±1.1 | 20.1±2.1 | 4.5±0.4 | 4.41±0.19 | 17.3±1.9 | 1.80±0.13 |
min. – max. | 17.8–30.7 | 7.9–11.9 | 16.8–23.8 | 3.8–5.2 | 3.77–4.89 | 14.0–20.9 | 1.42–2.16 |
cretica | n = 2 | n = 2 | n = 2 | n = 2 | n = 2 | n = 2 | n = 2 |
mean ± SD | n.a. | n.a. | n.a. | n.a. | n.a. | n.a. | n.a. |
min. – max. | 22.2–24.2 | 9.1–9.6 | 21.4–22.4 | 4.6–4.9 | 4.36–4.87 | 18.1–18.2 | 1.86–1.99 |
feri | n = 1 | n = 1 | n = 1 | n = 1 | n = 1 | n = 1 | n = 1 |
mean ± SD | n.a. | n.a. | n.a. | n.a. | n.a. | n.a. | n.a. |
min. – max. | 27.8 | 9.6 | 21.7 | 4.9 | 4.43 | 16.1 | 1.67 |
forcipata | n = 14 | n = 14 | n = 14 | n = 14 | n = 14 | n = 14 | n = 14 |
mean ± SD | 24.2±2.7 | 9.5±0.8 | 19.0±1.9 | 4.4±0.4 | 4.38±0.27 | 17.9±0.8 | 1.90±0.14 |
min. – max. | 21.6–30.8 | 8.3–11.00 | 16.6–22.8 | 4.0–5.3 | 3.96–5.04 | 16.7–19.6 | 1.63–2.12 |
francisae | n = 38 | n = 38 | n = 37 | n = 37 | n = 37 | n = 38 | n = 38 |
mean ± SD | 24.1±2.4 | 9.6±0.6 | 22.2±1.0 | 4.5±0.2 | 4.93±0.28 | 16.6±1.4 | 1.73±0.11 |
min. – max. | 19.5–31.5 | 8.5–10.5 | 20.9–24.9 | 4.1–5.1 | 4.55–5.62 | 13.8–19.3 | 1.52–1.99 |
gemellata | n = 2 | n = 2 | n = 2 | n = 2 | n = 2 | n = 2 | n = 2 |
mean ± SD | n.a. | n.a. | n.a. | n.a. | n.a. | n.a. | n.a. |
min. – max. | 20.5–22.1 | 7.4–10.0 | 16.5–22.4 | 4.2–4.6 | 3.93–4.92 | 13.8–14.2 | 1.42–1.86 |
giuliae | n = 30 | n = 30 | n = 30 | n = 30 | n = 30 | n = 30 | n = 30 |
mean ± SD | 24.2±2.7 | 10.3±0.4 | 22.3±0.9 | 4.8±0.2 | 4.65±0.19 | 17.7±1.5 | 1.73±0.15 |
min. – max. | 18.7–29.8 | 9.4–10.9 | 21.1–24.2 | 4.5–5.4 | 4.38–5.13 | 15.7–21.2 | 1.45–2.12 |
jacquelinae | n = 7 | n = 7 | n = 7 | n = 7 | n = 7 | n = 7 | n = 7 |
mean ± SD | 28.3±2.9 | 10.6±0.4 | 26.2±0.8 | 5.3±0.1 | 4.96±0.16 | 20.3±0.9 | 1.91±0.09 |
min. – max. | 24.9–33.6 | 10.2–11.1 | 25.2–27.5 | 5.1–5.4 | 4.74–5.29 | 19.4–21.8 | 1.76–2.04 |
latens | n = 8 | n = 8 | n = 9 | n = 9 | n = 9 | n = 8 | n = 8 |
mean ± SD | 20.3±2.3 | 98.7±0.5 | 18.8±1.9 | 4.3±0.3 | 4.40±0.25 | 14.9±0.8 | 1.72±0.05 |
min. – max. | 17.5–23.3 | 8.1–9.4 | 17.1–22.9 | 3.9–4.8 | 3.98–4.77 | 13.8–16.4 | 1.64–1.79 |
mariannae | n = 15 | n = 15 | n = 14 | n = 14 | n = 14 | n = 15 | n = 15 |
mean ± SD | 22.4±2.8 | 8.9±1.0 | 19.0±2.4 | 4.3±0.3 | 4.43±0.27 | 15.9±1.2 | 1.80±0.22 |
min. – max. | 19.1–27.5 | 7.5–10.3 | 16.5–22.8 | 3.9–4.9 | 3.92–4.74 | 13.6–17.5 | 1.41–2.23 |
marietheresae | n = 6 | n = 6 | n = 6 | n = 6 | n = 6 | n = 6 | n = 6 |
mean ± SD | 23.6±3.3 | 8.3±0.7 | 18.5±2.1 | 4.2±0.2 | 4.36±0.39 | 17.9±1.1 | 2.15±0.14 |
min. – max. | 20.5–28.8 | 7.4–9.5 | 16.6–22.6 | 3.9–4.4 | 3.95–5.16 | 15.9–18.9 | 1.98–2.38 |
pallipes | n = 1 | n = 1 | n = 1 | n = 1 | n = 1 | n = 1 | n = 1 |
mean ± SD | n.a. | n.a. | n.a. | n.a. | n.a. | n.a. | n.a. |
min. – max. | 21.7 | 9.5 | 23.8 | 4.6 | 5.23 | 13.5 | 1.43 |
smyrnensis | n = 5 | n = 5 | n = 5 | n = 5 | n = 5 | n = 5 | n = 5 |
mean ± SD | 20.0±1.8 | 9.9±0.3 | 23.6±0.9 | 4.6±0.1 | 5.18±0.18 | 17.6±0.8 | 1.78±0.12 |
min. – max. | 17.5–22.6 | 9.5–10.3 | 22.4–24.7 | 4.4–4.7 | 4.92–5.48 | 16.6–18.6 | 1.61–1.96 |
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.
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
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 (
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.
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.
Eupholidoptera gemellata Willemse & Kruseman, 1976: 136.
Morphological description.
The species was described after a single male was collected in 1973. Pitfall catches made in 2000–2001 at Mt. Psiloritis at 1950 m above Lochria and Agia Marina caught 11 males and 8 females. Opportunity is taken here to describe the female and illustrate important morphological structures with stacked images.
Holotype, 3 ♂, 2 ♀ (for details see Suppl. material
Frontal part of head (Fig.
Female. Examined specimens. 2 ♀: RETHIMNO: Psiloritis, above Lochria, FC1602 1♀
General appearance and size as male (Figs
See Tables
The song of this species has not yet been recorded.
Males differ from congenerics in the stout, straight cercus (Figs
The holotype was collected on Mt. Idi at 1650 m near the spring of Skaronero. Additional specimens collected in pitfall traps at a site northwest of Skaronero at 1950 m above Lochria between 15 September 2000 and 12 June 2001 (Fig.
Rocky mountain slopes with phrygana.
The holotype was collected at 1650 m on 28 July. The pitfalls that trapped the species were positioned at 1910 m and emptied on 15 September and 30 October 2000, and again on 12 June 2001.
Eupholidoptera giuliae Massa, 1999: 72.
Morphological description.
2 ♂, 1♀ (paratypes); 52 ♂, 30 ♀ (for details see Suppl. material
Frontal part of head (Fig.
See Tables
Based upon the sound recordings of 5 specimens (50 syllables measured), the song of E. giuliae – 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. giuliae, the syllable duration is ~ 199 ms, with a syllable rate up to ~ 1/s. There are no published descriptions of the song of this species. 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 giuliae see Suppl. material
Along the south coast and more to the northeast up toward the town of Rethimno males show little variation in cerci, anal tergite, subgenital plate or titillator. Styli are small, mostly pointing inward but in some specimens somewhat downward. In the titillator the two apical arms are mostly divergent but sometimes almost parallel and close to each other. It is unclear whether such variation is structural, the result of the drying up process after killing or the age of the specimen in number of days after the final moult. Toward the northwest, in the municipalities of Chania and Apokoronas, male subgenital plates (Fig.
Males differ from congenerics in the wide, upturned, spineless subgenital plate (Figs
The species was described from Chora Sfakion and a site 2.5 km east of Argoules along the southwestern coast of Crete (
Based on current data E. giuliae is a lowland species occurring from sea level up to some 500 m. It has been found in a variety of habitats ranging from very dry phrygana covered hills and Quercus forest, to rather wet lush vegetations including ferns.
Adults of this species have been collected by hand from 10 May to 24 June. This is also the period during which most pitfall catches were made but at least in one instance E. giuliae adults were caught in a trap that had been set 20 August. This indicates that although adults may appear early in the season and may be most numerous in June, they can still be found until late August.
Eupholidoptera jacquelinae Tilmans, 2002: 157.
Morphological description.
Holotype, allotype, 2 ♂ (paratypes); 4 ♂, 6 ♀ (for details see Suppl. material
Frontal part of head (Fig.
See Tables
Based upon the sound recordings of one specimen, the song of E. jacquelinae, 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. jacquelinae, the syllable duration is ~ 231 ms (Fig.
Habitus Eupholidoptera spp. in dorsal and lateral view 224, 225 Eupholidoptera astyla ♀ Mt. Idhi CT1987.046.03 226, 227 Eupholidoptera cretica ♀ Mt. Lefka Omalos FC17807
For the description of E. jacquelinae in 2002 only three males and one female from Gavdos were available. At present more specimens are at hand, also from the islet of Gavdopoula. We compared seven males (5 from Gavdos and 2 from Gavdopoula) and seven females (also 5 from Gavdos and 2 from Gavdopoula). The males, compared with the holotype, show no differences in colour, marking, last abdominal tergite, subgenital plate with styli, cercus, or titillator (from three males). The females, however, compared to the allotype, show some variation in marking and the form of the subgenital plate. Most of the females show more black markings on the lateral lobes of the pronotum than the allotype, but even then, less markings than in the males. A bit more than half of the females studied possesses a subgenital plate that is not longer than wide as in the allotype, but as long as wide or even a little bit wider than long. In one female from Gavdos and one of Gavdopoula the median groove of the subgenital plate fades away toward the basis. In a small number of the females the subgenital plate show faint transverse wrinkles.
Male E. jacquelinae is differentiated from all other (Cretan) Eupholidoptera by its uniquely shaped, strongly elongated apical lobes of the subgenital plate (Figs
Field images Eupholidoptera spp. Crete 240 Eupholidoptera annamariae ♂ Kata Zakros CT1995.0530 241 Eupholidoptera astyla ♂ Mt. Idhi CT2004.08.14 242 Eupholidoptera jacquelinae ♀ Gavdos 243 Eupholidoptera francisae sp. nov. ♂ paratype Andikithira CT2002.004.07 244 Eupholidoptera francisae sp. nov. ♀ paratype Andikithira CT2002.004.12 245 Eupholidoptera francisae ♂ paratype Marouliana RMNH5087052 246 Eupholidoptera francisae ♀ paratype Marouliana RMNH5014917.
Restricted to the islets of Gavdos and Gavdopoula, south of western Crete (Fig.
The habitats where the species was found consists of rocky ground with sparse vegetation of low trees (Pinus brutia), thorny shrubs and smaller plants as well as sand dunes with Pistacia, Juniperus, and Tamarix. The male specimens of E. jacquelinae were collected by hand on bushes of Erica manipuliflora, Pistacia, Tamarix, and Pinus but not on prickly bushes like Juniperus or Sarcopoterium spinosum, a spiny shrublet very common on Crete but much less so on Gavdos. The females however were found hiding under the low spiny shrubs of Euphorbia acanthothamnos. Trap catches on Gavdos and Gavdopoula recorded the species between 8 and 270 m.
Field images Eupholidoptera spp. Crete 247 Eupholidoptera giuliae ♂ Prasies RMNH5087051 248 Eupholidoptera giuliae ♀ Rethymnon RMNH5014921 249 Eupholidoptera latens ♂ Kolympari RMNH553681 250 Eupholidoptera mariannae ♂ Kalavros RMNH5014907 251 Eupholidoptera mariannae ♀ Ag. Ioannis RMNH5014906 252 Eupholidoptera pallipes ♀ below Pakhnes 253 Eupholidoptera smyrnensis ♂ Makrigiannis RMNH5087053.
The holotype male together with a paratype male was collected on 11 June at 50 m. The allotype female together with another paratype male was collected as nymph in the period 30 April to 2 May, becoming adult 25 May. Additional females were collected on 5 August. Specimens were found in traps emptied between mid-March and mid-November.
Eupholidoptera latens Willemse & Kruseman, 1976: 134.
Morphological description.
Bioacoustics.
Holotype, allotype, 4 ♂, 3f (paratypes); 12 ♂, 7 ♀ (for details see Suppl. material
Frons (Fig.
See Tables
Based upon the sound recordings of nine specimens (49 syllables measured), the song of E. latens, 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. latens, the syllable duration is ~ 228 ms, with a syllable rate up to ~ 1/s. Published records (
Eupholidoptera latens is restricted to the northern and central Chania region in western Crete. The cerci and anal tergite show little variation across its range. Titillators in males from the Rodopou peninsula (Figs
Males differ from congenerics in the stout apical arms of the titillator (Figs
The species was discovered in 1973 at high altitudes on Mt. Lefka, western Crete and published records from this species originated only from this mountain and its foothills (Lakki). Specimens collected between 2016 and 2019 indicate the species also occurs at low altitudes in the northern and northeastern parts of Chania region (Fig.
The species occupies habitats from sea level to alpine regions between 1600 and 1800 m on Mt. Lefka. It hides in low prickly shrublets in the phrygana while at lower altitudes it was also found on shrubs of blackberry (Rubus) or gorse (Ulex).
At low altitudes adults appear around mid-May, at mid-level elevations toward the end of May or early June whereas at high altitudes it may take to the second half of July before the first adults appear.
Eupholidoptera mariannae
Willemse & Heller, 2001:
Morphological description.
Bioacoustics.
Holotype, 1 ♂ (paratype); 8 ♂, 15 ♀ (for details see Suppl. material
Frons (Fig.
See Tables
Based upon the sound recordings of two specimens (20 syllables measured), the song of E. mariannae, 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. mariannae, the syllable duration is ~ 172 ms. In the present recordings, the syllable repetition rate is very low. Published records (
Apical arms of the titillator in most specimens are fused, in some they become apically somewhat separated. The central pits in the anal tergite may be more or less well developed and more or less densely haired.
Examined specimens. 15 ♀: LASITHI: Agios Ioannis –
General appearance and colouration as male (Figs
Cercus short, conical hardly tapering but for apical third which is distinctly narrower, tip pointed, slightly upturned in profile, straight in dorsal view, covered with pale short and long hairs.
Subgenital plate (Figs
Ovipositor almost straight, apically slightly upcurved, 1.4–2.2× longer than pronotum.
Males differ from congenerics in the stout, upturned, proximally flattened cercus, pointing outward (Figs
The species was described from southwestern Lasithi. Recent findings indicate its distribution area also includes western central Lasithi from the eastern slopes of Mt. Dikti eastward up to Kalavros in the north and Koutsouras in the south (Fig.
The altitudinal range of E. mariannae is considerable. It ranges from sea level where it was found near Koutsouras, to 1475 m, the highest altitude at which it was found on Mt. Thrypti. The type series was found inside open pine forest and groups of planted olive trees. Near Kalavros the species was found on open hill slopes with quite a dense vegetation of small shrubs interspersed with taller shrubs. Near Kavousi (Pacheia Amos) the species was trapped on a hill covered by phrygana. Around Malles and Anadoli individual males were heard singing on the branches of olive trees a few meters above the ground.
At low altitudes adults can be found already in early May, at higher altitudes starting from 300–700 m up to 1475 m, adults appear in June or July and have been caught until the end of September and mid-October.
Pitfall catches collected on Mt. Dikti at a site above the Limnakaro plateau trapped a total of 127 specimens of Eupholidoptera which at first glance were identified as E. forcipata. Closer examination however revealed differences with E. forcipata based on which a species new to science is described here.
Type locality. Greece, Crete, Lasithi, Dikti Mt., above Limnakaro plateau, SE of Ag. Anastasi mountain refuge, SW Spathi Madharas, 35.1107°N, 25.4779°E, 1715 m
Holotype ♂,
Paratypes. 1 ♂
(not examined). 40 ♂, 37 ♀ in alcohol (
Male. General appearance (Figs
Stridulatory file with 211–216 teeth (including proximal and distal ones), density of teeth in middle two thirds of the file 37–42 teeth per mm.
Anal tergite (Figs
Cercus (Figs
Subgenital plate (Figs
Titillator (Figs
General colouration (based on specimens kept in alcohol) yellowish brown. Head with the frontal part below antennae and eyes pale with two black dots (Fig.
Female. General appearance (Figs
See Tables
The song of this species has not yet been recorded.
Males differ from congenerics in the pointed back- and downwardly extended widened lobes of the anal tergite (Figs
Only known from a single location on Mt. Dikti above the Limnakaro plateau where the species was trapped in pitfall traps (Fig.
Mountain slopes at 1700 m in phrygana vegetation.
Pitfall traps in which the species was found were checked irregularly. Based on the three catching periods, adults can be found prior to early August up to at least early October.
The species is named in honour of Marie-Therèse Willemse-Dresen (1929–2017) wife and lifelong companion of Fer Willemse who contributed a large part of his entomological career to the study of the Orthoptera fauna of Greece, describing 40 species new to science from Greece, including four Cretan species of Eupholidoptera. In his last, and most challenging, paper on Chorthippus (Glyptobothrus) from Greece (
“This publication would not have been possible without help from my family. My wife’s patience and tolerance to my single-minded enthusiasm was almost boundless. Both the long hours spent during our travels in seemingly dull and uninteresting areas, as well as at home recording and studying have been accepted without much ado. For that I owe her an enormous amount of gratitude.”
It is in this spirit that we pay a tribute to Marie-Therèse. The fact that E. marietheresae sp. nov. is found on the same mountain and in the vicinity of E. feri, a species named after Fer Willemse, is making it even more appropriate.
Eupholidoptera pallipes Willemse & Kruseman, 1976: 135.
Morphological description.
Holotype, allotype, 5 ♂ (paratypes) (for details see Suppl. material
Frontal part of head (Fig.
See Tables
The song of this species has not yet been recorded.
Males differ from congenerics in the stout, straight cerci (Figs
The type series was collected in 1973 on Mt. Lefka at the saddle of Linoseli above Xyloskalo between 1600 m and 1800 m. Additional specimens were collected in pitfall traps operated in the summer of 1991 more to the east on Mt. Lefka above Limnia (Fig.
Rocky mountain slopes with phrygana between 1600 and 2440 m.
The type series was collected 5 August 1973. Pitfalls above Limnia trapped adults between early August and early September and during the entire month of October whereas a trap operated between early June and early July only contained nymphs.
Thamnotrizon smyrnensis Brunner von Wattenwyl, 1882: 336.
Olynthoscelis smyrnensis (Brunner von Wattenwyl, 1882): Bolivar 1899: 601.
Pholidoptera smyrnensis (Brunner von Wattenwyl, 1882): Ebner 1919: 157.
Eupholidoptera smyrnensis
(Brunner von Wattenwyl, 1882):
Morphological description.
Bioacoustics.
9 ♂, 5 ♀ (for details see Suppl. material
Frontal part of head (Fig.
See Tables
Based upon the sound recordings of one specimen (10 syllables measured) from Crete, the song of E. smyrnensis, 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. smyrnensis, the syllable duration is ~ 40 ms, the shortest in Cretan Eupholidoptera. In the present recordings, the syllable repetition rate is 2/s at maximum. The song is also characterised by syllables that are produced in compact series of 3–10s followed by a longer silence after which another series follows. The first few syllables in a series are weaker than the following ones. This pattern has so far not been found in other species of Eupholidoptera in Crete. Recordings from other Greek Islands (Rhodes and Naxos) and published by
Males differ from congenerics in the stout, inward curved cerci (Figs
Eupholidoptera smyrnensis is one of the most widespread species of the genus. Its range covers western Anatolia and the southern Balkan (southern Bulgaria, southeastern Republic of North Macedonia, and north-western Greece) (
Although on the Greek mainland E. smyrnensis is found up to 1200 m, in Crete, it has only been found between 25 m and 340 m. Unlike most other Eupholidoptera species in Crete, E. smyrnensis is not found in spiny shrubs on the ground but lives in taller spiny bushes like blackberries (Rubus).
Based on hand and pitfall catches E. smyrnensis becomes adult in June and near Limni Kourna adults have still been trapped after 20 August, probably being active well into September and possibly October.
The aligned concatenated dataset, which consisted of 1684 bp including 375 variable and 292 parsimony informative sites, involved ten ingroup and two outgroup taxa (respectively 36 ingroup and two outgroup haplotypes). The NADH2 fragment consisted of 855 bp including 353 variable and 279 parsimony informative sites, and the ITS had 829 bp with gaps, with 22 variable and 13 parsimony-informative sites. No numt signs were detected in alignments of protein-coding sequences, and the saturation tests did not show signs of significant saturation. Best substitution models for the partitioned dataset were as follows: NADH2, positions 1 and 2 – gamma, position 3 – gamma+ invariable sites; ITS – proportion of invariable sites.
Our phylogenetic analysis provided well resolved phylogeny of the studied taxa, showing strong support for all nodes (Fig.
The taxa E. latens, E. giuliae, E. francisae, and E. astyla form a monophyletic lineage that splits into two major clades. The first clade contains all specimens occurring in western and southwestern Chania and Andikithira, showing very low genetic distances to each other (see also Fig.
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 1684bp concatenated alignment of the NADH2 + ITS fragments. Node values at branches show node support with BI posterior probabilities. Different colours correspond to the different morphological species. See Suppl. material
Some Cretan species appear to be paraphyletic: E. marietheresae is nested within E. mariannae despite their large morphological differences; an E. annamariae specimen found at the boundary of the E. annamariae and E. mariannae distributions appears to be more closely related to E. mariannae than to E. annamariae, despite its distinct morphology; and an E. astyla specimen is more closely related to E. giuliae than to other E. astyla’s.
The Eupholidoptera giuliae-latens-francisae complex: geographic distribution of five morphological entities in the Eupholidoptera giuliae-latens-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).
The position of E. marietheresae inside the E. mariannae clade is an interesting observation, given their distinct morphological differences. Additional study is required, including the incorporation of E. forcipata sequences in the phylogenetic tree (the species morphologically closest to E. marietheresae), of more E. marietheresae specimens, and additional molecular (mostly nuclear) markers, in order to decipher the relationship between these species. Here it should be mentioned that the molecular results from
Eupholidoptera jacquelinae, the endemic of the island of Gavdos, is the basalmost taxon of the Cretan clade. The only non-endemic Eupholidoptera on Crete, E. smyrnensis, is nested firmly within conspecifics from Anatolia. It should be noted that the nuclear fragment (ITS) showed very low interspecific variability. As a result, the topography of the concatenated tree is mostly influenced by the mitochondrial gene NADH2.
Together with western and southern Anatolia, Crete is a biodiversity hotspot for Eupholidoptera (
The current study revealed that based on morphological traits E. giuliae, E. latens and E. francisae may share a complex phylogeographical relationship.
Based on the shape of the anal tergite, subgenital plate, stylus and titillator of the male, five subgroups can be distinguished in western Crete and Andikithira (Table
Morphological differences across the E. latens-E. giuliae-E. francisae complex (see Fig.
Subgroups | Anal tergite excision-teeth | Subgenital plate | Styli | Titillator |
---|---|---|---|---|
1. Eastern Chania + western Rethimni | wide-short | compact | short-inward | compact – long hooks |
2. Northeastern Chania incl. Akrotiri | narrow-long | intermediate | long-downward | slender – long hooks |
3. Northern and central Chania | wide-short | slender | long-downward | compact – short hooks |
4. Western and southwestern Chania | narrow-short | very slender | short to long, downward | compact – short hooks |
5. Andikithira | narrow-short | extremely slender | short, downward | compact – short hooks |
Following the above pattern, populations in the western and southwestern corner of Crete (subgroup 4) represent a genetically well-outlined lineage, sharing the nuclear ITS fragment and its mitochondrial genome with the population on Andikithira (subgroup 5) (genetic distances of the nuclear internal transcribed spacers show very low variation among Eupholidoptera). The two islands have been isolated since the beginning of the Pleistocene (2.6 Mya), with only a narrow length of sea separating them during the Last Glacial Maximum (
Based on the molecular results and the low morphological differences between populations from Andikithira and western and southwestern Chania (subgroups 4 and 5 in Table
The second clade may be regarded as a monophyletic mitochondrially-defined species complex of three subclades – the typical E. giuliae (morph-group 1), a clade formed by morph-groups 2 and 3, and a clade formed by E. astyla samples. The very low genetic mitochondrial distances between morphologically well outlined taxa (i.e., E. astyla and the rest) may point to former population crises (bottlenecks), where mitochondrial genome was shared between two or more taxa. At the same time, genetic drift caused by genetic bottlenecks may have contributed to unique characteristics (for instance, the peculiar shape of the titillators in E. astyla). On the other hand, the existence of specimens with symmetrical titillators and the intermixture of a tentatively identified female of E. astyla within the E. giuliae s.str. subclade (Fig.
An alternative scenario of the systematics of this western species complex may be proposed. Since the clade of morph-groups 2 and 3 show intermediate and variable morphology between specimens of E. francisae (morph-group 4) and E. giuliae (morph-group 1), and they intermix at the phylogenetic tree, those populations may represent a hybridogenic lineage of two formerly partially speciated taxa or may still be in the ‘grey zone’ of their evolutionary differentiation where they can either become distinct taxa or merge back into a common genetic pool in future.
The above scenario may not be unique and could be expected in other sibling species of Cretan Eupholidoptera that express intermediate or variable morphological characters (e.g., E. mariannae and E. annamariae). As results based on morphology and genetics are not unequivocal and boundaries do not match, as yet no taxonomic decision has been taken for morph-groups 1, 2, and 3. For the time being, both the eastern groups (morph-groups 1 and 2) have been assigned to E. giuliae, whereas the central group (morph-group 3) matches E. latens.
The diversity of male insect genitalia is well known (
Although in some orders of insects, asymmetry of genitalia is the ground plan, in most insects, including Orthoptera, it is very rare (
The song of Eupholidoptera is quite uniform throughout the genus and is even considered a not very reliable character for discerning species (
In quite some recordings it is not easy to clearly discern the opening hemisyllable, leaving discussion whether it is just weak and may be not well recorded or it is absent, with no sound produced during this wing movement. As a main character of the species’ song may be the duration of the syllable, the absence of the opening hemisyllable in some recordings or specimens accounts for inaccurate measurements of this song character. The same holds for the effect of temperature on the duration of syllables, which may last twice as long with temperatures differing only as little as 2–5 °C. This adds to the fact that in many cases not so many specimens and not so much time (actively) singing has been available to us to compare the song of the species thoroughly. Bioacoustic measurements presented in Fig.
Most Cretan species of Eupholidoptera are allopatric and a female will only hear the calling song of conspecific males. The syllable duration between most Cretan species is largely overlapping and not distinctive (but long for Eupholidoptera in general;
Trap no. | Coll. date | Location | GPS Coordinates | Alt. (m) | Species 1 | Species 2 |
---|---|---|---|---|---|---|
FC111 | 30/10/1996 | Limni Kourna | 35.3269, 24.2790 | 25 | giuliae | smyrnensis |
FC495 | 10/07/1997 | Limni Kourna | 35.3269, 24.2790 | 25 | giuliae | smyrnensis |
FC70 | 20/08/1996 | Limni Kourna | 35.3269, 24.2790 | 25 | giuliae | smyrnensis |
FC1536 | 05/08/2000 | Dikti Mt. | 35.1107, 25.4779 | 1715 | astyla | marietheresae sp. nov. |
FC1606 | 02/10/2000 | Dikti Mt. | 35.1107, 25.4779 | 1715 | astyla | marietheresae sp. nov. |
FC1655 | 09/01/2001 | Dikti Mt. | 35.1107, 25.4779 | 1715 | astyla | marietheresae sp. nov. |
FC1602 | 15/09/2000 | Idi Mt. | 35.1973, 24.7920 | 1910 | forcipata | gemellata |
FC1651 | 30/10/2000 | Idi Mt. | 35.1973, 24.7920 | 1910 | forcipata | gemellata |
FC1916 | 12/06/2001 | Idi Mt. | 35.1973, 24.7920 | 1910 | forcipata | gemellata |
FC17807 | 19/10/2018 | Lefka Mt. | 35.3524, 23.9050 | 1200 | cretica | latens |
Findings presented here, collected over the past 30 years, show that the known distribution range for five Eupholidoptera species in Crete (E. annamariae, E. astyla, E. giuliae, E. latens, and E. mariannae) is larger than previously known (Fig.
Of the 14 species of Eupholidoptera treated in this paper, six species have only been found at higher altitudes (above 1000 m): E. pallipes, E. cretica, E. gemellata, E. forcipata, E. marietheresae sp. nov., and E. feri, five species are restricted to lower altitudes (below ca. 500 m): E. francisae sp. nov., E. giuliae, E. jacquelinae, E. annamariae, and E. smyrnensis, whereas data presented here indicate that E. astyla, E. latens, and E. mariannae are found across a wide altitudinal range from sea level to higher altitudes.
Pitfall traps are widely used to monitor ground dwelling invertebrates. For Orthoptera trapping results provide a contradictory picture (
Little information was available on the sympatric occurrences of different Eupholidoptera species. The syntopic occurrence of latens/pallipes on Mt. Lefka and gemellata/forcipata on Mt. Idi was conjected but not proven.
However, one must bear in mind that the period between activating the traps and collecting the accumulated specimens was at least two months. Therefore, it cannot be ruled out that despite the sympatry, species do not (completely) overlap in their phenology or daily activity patterns. Interestingly, in nine of the ten trapping events a species with unusual song pattern is involved, either smyrnensis with shorter syllables than all others or forcipata (respectively the similar marietheresae sp. nov. with probably similar song) with distinctly longer syllables than all others (see Fig.
Recently, a Red List for all European Orthoptera species has been compiled, including the Eupholidoptera species of Crete (
Current IUCN Red List Status for Eupholidoptera species of Crete and adjacent islands.
Species | Common name | IUCN RLA status* |
---|---|---|
E. annamariae | Annamaria’s Marbled Bush-cricket | VU |
E. astyla | Mount Ida Marbled Bush-cricket | EN |
E. cretica | Cretan Marbled Bush-cricket | VU |
E. feri | Fer’s Marbled Bush-cricket | CR |
E. forcipata | Idi Marbled Bush-cricket | VU |
E. francisae sp. nov. | Francis’s Marbled Bush-cricket | Not Evaluated |
E. gemellata | Skaronero Marbled Bush-cricket | VU |
E. giuliae | Giulia’s Marbled Bush-cricket | VU |
E. jacquelinae | Jacqueline’s Marbled Bush-cricket | VU |
E. latens | Hidden Marbled Bush-cricket | VU |
E. mariannae | Marianne’s Marbled Bush-cricket | VU |
E. marietheresae sp. nov. | Marietherese’s Marbled Bush-cricket | Not Evaluated |
E. pallipes | Pale-legged Marbled Bush-cricket | VU |
E. smyrnensis | Smyrna Marbled Bush-cricket | LC |
To further close the gaps in our knowledge, observations made during excursions by tourists in Crete can be very useful especially if these are uploaded to online platforms like iNaturalist and Observation.org. This paper includes nine observations on Eupholidoptera from Crete (Suppl. material
Fourteen species of Eupholidoptera in the Cretan area is a remarkable gathering of taxa, albeit not unique across various invertebrate orders. Other examples include for instance Dendarus (Coleoptera: Tenebrionidae) with 13 species (
This high concentration of endemic species of Eupholidoptera, and especially the presence of some primitive representatives (E. gemellata, E. pallipes), has led previous researchers to propose the southeastern Aegean plate, possibly including Crete, as the origin place of the last common ancestor of Eupholidoptera (
Of the fourteen species of Eupholidoptera only one, E. smyrnensis, has a large distribution area. The finding of Eupholidoptera smyrnensis on Crete adds new information to the phylogeography of the Eupholidoptera chabrieri group of species as presented by
Only one Eupholidoptera species is not found on Crete itself but restricted to surrounding islands. Gavdos and the small islet of Gavdopoula are home to E. jacquelinae. The islands are surrounded by a fairly extensive shelf that was exposed above sea level during Mio-Pliocene but were never connected to Crete during the Quaternary (2.58 Mya up to now) (
Another interesting phylogeographical matter is that of Eupholidoptera francisae occurring on the island of Andikithira (32 km NW of western Crete) and in the extreme western/southwestern part of Crete. The faunal affinities between Andikithira and W. Crete and their biogeographical implications in the area, have a long story of theories and debates, starting from “Boettger’s line” (between Kithira and Andikithira) and the Peloponnese-Kithira-Andikithira-Crete relations (
One more important question is, what caused the diversification of the 11 endemic Eupholidoptera species restricted to Crete itself and what factors contributed to their current distribution patterns? Although phylogeographic studies over the past 30 years revealed two major geological events that contributed to the diversification in the Aegean – the formation of the mid-Aegean trench (MAT) at the end of the Miocene (12–9 Mya) that separated Crete from the west Aegean and Anatolia, and the final isolation of Crete from Peloponnisos after the Messinian Salinity Crisis (MSC, 5.33 Mya) – solid evidence of exact patterns of diversification via in situ speciation within individual islands in the Aegean, i.e. Crete, remains still scarce (
Several studies on monophyletic lineages of invertebrates and vertebrates (Table
Cretan invertebrate and vertebrate monophyletic lineages and time of divergence of East-West clades.
Taxon | Order | Number of subclades | Diverged approximately at | Reference |
---|---|---|---|---|
Cyrtocarenum cunicularium | Araneae | 2 | 3.3 Mya |
|
Poecilimon cretensis | Orthoptera | 2 | 0.6 – 1.2 Mya |
|
Thaumetopoea wilkinsoni | Lepidoptera | 2 | < 1 Mya |
|
Reticulitermes | Blattodea | 4 | < 1 Mya | Fig. |
Carabus banoni | Coleoptera | 2 | 1.47 Mya | Vlachopoulos et al. (pers. Comm.) |
Podarcis cretensis | Squamata | 2 | 5.42 – 4.36 Mya |
|
East-west divergences are also evident in Cretan Eupholidoptera lineages, but the question whether the species found in Crete and the adjacent islands are polyphyletic as concluded by
An example of another polyphyletic genus under investigation on Crete is Dendarus (Coleoptera: Tenebrionidae) that consists of three distinct lineages on the island (two mitochondrial, one nuclear gene locus, and 61 morphological characters;
We are thankful to the Directorate General for the Protection and Development of Forests and Rural Environments of the Ministry for Environment and Energy in Greece for issuing permits to Luc Willemse and Jos Tilmans in 2017 (154383/852) and 2019 (178527/93/22-1-2-19) to collect Eupholidoptera for further studies.
Luc Willemse wishes to express his gratitude to Panoraia Zacharopoulou who located many specimens during their 2017 trip to Crete.
Jos Tilmans thanks his wife Jacqueline Tilmans-Smid for her never-ending help and endurance in collecting Eupholidoptera specimens during the several trips to Crete, Gavdos, and Andikithira between 1987 and 2019.
Dragan Chobanov would like to thank the grants supporting the Orthoptera Species File and the Greek authorities (permission 154812/951 from 6 June 2017).
Nefeli Kotitsa is grateful to the Linné Systematics Research Fund and the Erasmus+ Traineeship program. She and Apostolos Trichas would also like to give huge thanks to the researchers and students of the National History Museum Crete and the University of Crete that carefully and methodically collected, sorted, and stored the Eupholidoptera specimens during the trapping programs between 1987 and 2019.
Klaus-Gerhard Heller would like to especially thank Martina Heller and Marianne Volleth for their diverse help during their travels in Crete. Without the help of Martina we would have not even noticed the animals during the trip in 2016.
Roy Kleukers, Alexandros Quartarone, and Peter Landert are thanked for allowing us to use their photographs of live animals and Damien Sevrin for his image and record of Eupholidoptera. Juergen Deckert and Claudia Hömberg are gratefully acknowledged for the loan of the E. astyla type specimens described by Ramme. Laboratory staff of Naturalis Biodiversity Center, in particular Frank Stokvis and Roland Butot, are thanked for their assistance with the stacking microscope equipment.
For helpful comments we are grateful to the reviewers Axel Hochkirch and Sigfrid Ingrisch.
List of localities, specimens, and repositories of Eupholidoptera from Crete, Gavdos, Gavdopoula, and Andikithira
Data type: occurrence data (excel document)
Explanation note: The table contains all published and unpublished localities, specimens collected and their repositories for all Eupholidoptera species from Crete and its satellite islands known to the authors.
Examined specimens
Data type: specimen studied (excel document)
Explanation note: This table lists examined specimens with an indication of how they are stored (D: dry pinned; W: in alcohol), whether they have been measured (mm), photographed (image), sampled for DNA analysis (DNA), or audio recorded (audio). Abbreviations used: √: done; S: stacked images(s); F: field image from life specimen; H: habitus image from collection specimen; FC: trap. For more details of location and trap see Suppl. material
Sound recordings and song data
Data type: bioacoustic recordings (excel document)
Explanation note: table with detailed information on sound recordings and song data. The tab sound recordings summarises details of sound recordings of Eupholidoptera species from Crete, Andikithira, and Gavdos. Recordings are in the collection of Jos Tilmans, Baudewijn Odé, and Klaus-Gerhard Heller. The tab song details provides information about the ambient temperature and mean syllable duration for individual specimen.
Details of specimens used in phylogenetic analysis
Data type: table (excel document)
Explanation note: Details of specimens used for the Bayesian inference phylogenetic tree of 9 Eupholidoptera species from Crete, Andikithira, and Gavdos (see Figs
NADH2___ITIS_sequences
Data type: table (excel document)