Corresponding author: Luc Willemse (
Academic editor: Zhu-Qing He
Being nocturnal, hiding in prickly bushes and shrubs during the day,
Willemse L, Tilmans J, Kotitsa N, Trichas A, Heller K-G, Chobanov D, Odé B (2023) A review of
Occurrence records
Published record of
Species | Number of locations | Altitude (m) | Reference |
---|---|---|---|
|
2 | 50–200 | |
|
10 | 800–1850 | |
|
1 | not indicated |
|
|
1 | 1100 |
|
|
3 | 1700–1850 | |
|
1 | 1650 |
|
|
3 | 0–175 | |
|
1 | 50 | |
|
7 | 500–1800 | |
|
4(1) | 500–700 | |
|
1 | 1600–1800 |
|
1: includes two locations of paratypes described by
Hand catches included scanning larger shrubs (
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
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.
Traps
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
Sequence data for one protein-coding mitochondrial gene (NADH dehydrogenase subunit 2 –
Polymerase chain reactions were performed using Thermo Scientific DreamTaq Hot Start Master Mix according to the manufacturer’s instructions. Temperature cycling for the
Additional sequences for both loci were obtained from 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 (
Coordinates are presented in decimal degrees (
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
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 (
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
Measurements
A list summarising all known localities, specimens, and repositories is presented in Suppl. material
Table
Colour pattern abdomen from above
Published and unpublished records (period 1987–2020) of
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 | |
---|---|---|---|---|---|---|---|---|---|---|---|
|
5–550 | 2 | 5 | 3 | 0 | 0 | 0 | 2 | 2 | 0 | 14 |
|
5–1850 | 10 | 6 | 24 | 0 | 0 | 1 | 3 | 1 | 0 | 45 |
|
1165–1234 | 1 | 0 | 4 | 0 | 0 | 0 | 0 | 0 | 0 | 5 |
|
1100 | 1 | 0 | 0 | 0 | 0 | 0 | 0 | 0 | 0 | 1 |
|
1350–2225 | 3 | 2 | 3 | 0 | 0 | 0 | 0 | 0 | 0 | 8 |
1–835 | 0 | 2 | 1 | 0 | 0 | 0 | 2 | 2 | 9 | 16 | |
|
1650–1910 | 1 | 0 | 1 | 0 | 0 | 0 | 0 | 0 | 0 | 2 |
|
0–525 | 3 | 4 | 9 | 0 | 7 | 1 | 3 | 1 | 4 | 32 |
|
30–270 | 1 | 0 | 8 | 0 | 1 | 0 | 0 | 0 | 0 | 10 |
|
20–1815 | 7 | 3 | 4 | 1 | 0 | 1 | 1 | 2 | 2 | 21 |
|
0–1475 | 4 | 2 | 9 | 0 | 0 | 0 | 2 | 0 | 0 | 17 |
1715 | 0 | 0 | 1 | 0 | 0 | 0 | 0 | 0 | 0 | 1 | |
|
1600–2440 | 1 | 0 | 1 | 0 | 1 | 0 | 0 | 0 | 0 | 3 |
|
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. |
|
– | Frons with isolated dark spots or black crossband (Figs |
|
2 | Females |
|
– | Males |
|
3 | Subgenital plate with pair of elongate concavities divided by median ridge, apical lobes touching (Fig. |
|
– | Subgenital plate convex or proximally concave or with pits, apical lobes separated (Figs |
|
4 | Gavdos or Gavdopoula |
|
– | Crete or Andikithira |
|
5 | Subgenital plate wider to much wider than long (Figs |
|
– | Subgenital plate as wide as long or elongated (Figs |
|
6 | Hind margin of subgenital plate with wide and deep median excision (Fig. |
|
– | Hind margin of subgenital plate differently formed (Figs |
|
7 | Hind margin of subgenital plate medially concave, without excision (Fig. |
|
– | Hind margin of subgenital plate with distinct excision (Figs |
|
8 | Subgenital plate with apical lobes rounded (Fig. |
|
– | Subgenital plate with apical lobes pointed (Figs |
|
9 | Subgenital plate 2.0× wider than long, proximal pits small, widely separated (Fig. |
|
– | Subgenital plate 1.4× wider than long, proximal pits large, closer to each other (Fig. |
|
10 | Front of head with large black patches (Fig. |
|
– | Front of head with black dots (Figs |
|
11 | Apex subgenital plate in profile (measured in a straight line parallel to the ovipositor) not reaching or surpassing proximal half of gonangulum (Fig. |
|
– | Apex subgenital plate in profile (measured in a straight line parallel to the ovipositor) reaching or surpassing distal half of gonangulum (Fig. |
|
12 | Subgenital plate proximally convex, flattened or slightly concave (Fig. |
|
– | Subgenital plate proximally with one wide or two separate pits (Figs |
|
13 | Apical lobes of subgenital plate not well produced, rectangular (Fig. |
|
– | Apical lobes of subgenital plate more produced, rectangular to acute (Figs |
|
14 | Subgenital plate proximally with two distinct pits separated by a median keel (Fig. |
|
– | Subgenital plate proximally with a single or two pits (Figs |
|
– | (Mt. Dikti, Katharo plains) |
|
15 | Styli pointing backwards (Figs |
|
– | Styli pointing downward or inward (Figs |
|
16 | Cercus with subbasal side tooth (Figs |
|
– | Cercus unarmed (Figs |
|
17 | Pronotum pale (Fig. |
|
– | Pronotum with black band or patches (Fig. |
|
18 | Styli very long, 5–6× longer than wide (Figs |
|
– | Styli short, 2× longer than wide (Figs |
|
19 | Subgenital plate with long spine at base of stylus (Fig. |
|
– | Subgenital plate without a spine at base of stylus (Fig. |
|
20 | Hind margin of anal tergite from the cercus downward straight, side flaps gradually narrowing (Fig. |
|
– | Hind margin of anal tergite from the cercus downward S-curved, side flaps first widening before narrowing (Fig. |
|
21 | Anal tergite distally extended into two very long, spined hooks pointing downward (Fig. |
|
– | Anal tergite distally extended into short pointed lobes (Figs |
|
22 | Pointed lobes of anal tergite close together (Figs |
|
– | Pointed lobes of anal tergite widely separated (Figs |
|
23 | Cercus unarmed (Fig. |
|
– | Cercus with subbasal side tooth (Figs |
|
24 | Cercus conical (Fig. |
|
– | Cercus flattened and widened proximally (Fig. |
|
25 | Subgenital plate as wide as long (Fig. |
|
– | Subgenital plate elongate, tapering toward the apex (Figs |
|
26 | Tips apical lobes subgenital plate with a tooth (Figs |
|
– | Tips apical lobes subgenital plate always without a tooth (Figs |
|
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.
1 ♂, 1 ♀ (
Frontal part of head (Fig.
Colour pattern of head, frontal view
Colour pattern of pronotum, dorsal view
Female subgenital plate in ventral view
Female subgenital plate in lateral view
Female subgenital plate in lateral view
See Tables
Based upon the sound recordings of 6 specimens (53 syllables measured), the song of
The colour pattern and genitalia in
Males differ from congenerics in the anal tergite (Figs
Male anal tergite in dorsal view
Male anal tergite in caudal view
Male anal tergite in lateral view
Male cercus in dorsal view
Male cercus in lateral view
Male subgenital plate in ventral view
Male subgenital plate in lateral view
Male stylus in lateral view
Male titillator in ventral view
Male titillator in lateral view
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.
1 ♂ (
Frontal part of head (Fig.
In 1927
11 ♀: RETHIMNO: Idhi Mt., Idhaio Andro -1987.041.02 (
General appearance (Figs
See Tables
Based upon the sound recordings of two specimens (20 syllables measured), the song of
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
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
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.
1 ♂, 2 ♀ (for details see Suppl. material
Frontal part of head (Fig.
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
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.
Frontal part of head (Fig.
See Tables
Based upon the sound recordings of 1 specimen (10 syllables measured), the song of
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
The Katharo plain lies at an altitude of 1100 m. The type specimens were collected in late August.
Frontal part of head (Fig.
See Tables
Based upon the sound recordings of one specimen (30 syllables measured), the song of
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.,
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
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
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
See Tables
Based upon the sound recordings of 15 specimens (153 syllables), the song of
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
Tip of male subgenital plate in ventral view
Measurements (in mm) and biometrics of male
Males |
|
|
||
---|---|---|---|---|
length body | ||||
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 | ||||
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 | ||||
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 | ||||
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 | ||||
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 | ||||
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 | ||||
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 | ||||
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 | ||||
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
|
|
|
|
|
length body | ||||
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 | ||||
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 | ||||
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. | ||||
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 | ||||
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 | ||||
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 | ||||
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 | ||||
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 | ||||
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 | ||||
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 | ||||
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
The male subgenital plate of
The females of
Oscillograms of
Diagnostic characters and character-states for
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 | ||||||||||||||||||||||
|
1 | 3 | 1 | 2 | 2 | 2 | 2 | 1 | 1 | 2 | 1 | 1 | 1 | 1 | 2 | 1 | 1 | 1 | 1–3 | 2 | 1 | 1 |
|
1 | 3 | 1 | 1 | 1 | 1 | 3 | 1 | 1 | 1 | 1 | 1 | 1 | 1 | 3 | 1 | 3 | 1 | 3–4 | 2 | 1 | 1 |
|
1 | 1 | 1 | 1 | 3 | 3 | 2 | 1 | 1 | 2 | 1 | 1 | 3 | 2 | 1 | 1 | 2 | 2 | 1 | 3 | 2 | 1 |
|
1 | 3 | 1 | 1 | 1 | 1 | 2 | 3 | 1 | 1 | 1 | 1 | 1 | 1 | 2 | 1 | 2 | 1 | 4 | 2 | 1 | 2 |
|
1 | 1 | 1 | 2 | 3 | 1 | 1 | 1 | 1–2 | 3 | 1 | 1 | 2 | 2 | 1 | 1 | 3 | 1 | 1 | 1 | 2 | 3 |
|
1 | 2 | 1 | 1 | 2 | 1 | 3 | 1 | 3 | 2 | 1 | 1 | 1 | 1 | 1 | 1 | 3 | 2 | 1–3 | 3 | 2 | 2–3 |
|
3 | 3 | 3 | 1 | 1 | 2 | 2 | 3 | 1–3 | 2 | 2 | 2 | 2 | 2 | 1 | 2 | 2 | 2 | 1 | 1 | 2 | 1 |
|
1 | 2 | 1 | 1 | 2 | 1 | 3 | 1 | 1–3 | 1 | 1 | 1 | 1 | 3 | 1 | 1 | 3 | 2 | 2–4 | 3 | 1 | 3 |
|
1 | 1 | 1 | 2 | 2 | 1 | 3 | 1 | 3 | 3 | 2 | 1 | 3 | 2 | 1 | 1 | 3 | 2 | 3–4 | 3 | 1 | 1 |
|
1 | 2 | 1 | 1 | 2 | 1 | 3 | 1 | (1)-3 | 2 | 1 | 1 | 2 | 1 | 1 | 1 | 3 | 2 | 2 | 3 | 1 | 2–3 |
|
1 | 3 | 1 | 1 | 1 | 3 | 1 | 3 | 2 | 1 | 1 | 1 | 1 | 1 | 2 | 1 | 3 | 1 | 3–4 | 2 | 1 | 2–3 |
|
3 | 1 | 1 | 2 | 3 | 1 | 1 | 1 | 1 | 3 | 1 | 1 | 2 | 2 | 1 | 1 | 3 | 1 | 1 | 2 | 2 | 3 |
|
3 | 3 | 3 | 1 | 1 | 2 | 2 | 3 | 2 | 2 | 2 | 2 | 2 | 2 | 1 | 2 | 1 | 2 | 2 | 1 | 2 | 1 |
|
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. |
|||||||||||||||||||||
b. pronotal disc | 1. pale (Fig. |
|||||||||||||||||||||
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 |
Measurements (in mm), ratios, and biometrics of male
Males | Body length | Pronotum length | Hind femur length | Hind femur width | Ratio length-width hind femur | Number of teeth |
---|---|---|---|---|---|---|
|
||||||
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 |
|
||||||
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 |
|
||||||
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 |
|
||||||
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 |
|
||||||
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 |
|
||||||
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 |
|
||||||
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 |
|
||||||
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 |
|
||||||
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 |
|
||||||
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 |
|
||||||
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 |
|
||||||
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 |
|
||||||
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 |
|
||||||
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 |
Measurements (in mm), ratios, and biometrics of female
Females | Body length | Pronotum length | Hind femur length | Hind femur width | Ratio length-width hind femur | Ovipositor length | Ratio length ovipositor-length pronotum |
---|---|---|---|---|---|---|---|
|
|||||||
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 |
|
|||||||
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 |
|
|||||||
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 |
|
|||||||
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 |
|
|||||||
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 |
|
|||||||
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 |
|
|||||||
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 |
|
|||||||
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 |
|
|||||||
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 |
|
|||||||
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 |
|
|||||||
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 |
|
|||||||
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 |
|
|||||||
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 |
|
|||||||
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
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
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
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
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.
Frontal part of head (Fig.
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.
2 ♂, 1♀ (
Frontal part of head (Fig.
See Tables
Based upon the sound recordings of 5 specimens (50 syllables measured), the song of
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
Male terminalia
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
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
Frontal part of head (Fig.
See Tables
Based upon the sound recordings of one specimen, the song of
Habitus
Habitus
For the description of
Male
Field images
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 (
Field images
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.
Frons (Fig.
See Tables
Based upon the sound recordings of nine specimens (49 syllables measured), the song of
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 (
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.
Distribution maps
Frons (Fig.
See Tables
Based upon the sound recordings of two specimens (20 syllables measured), the song of
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
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
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.
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
“
It is in this spirit that we pay a tribute to Marie-Therèse. The fact that
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.
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
Males differ from congenerics in the stout, inward curved cerci (Figs
Although on the Greek mainland
Based on hand and pitfall catches
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
Our phylogenetic analysis provided well resolved phylogeny of the studied taxa, showing strong support for all nodes (Fig.
The taxa
Bayesian inference phylogenetic tree of
Some Cretan species appear to be paraphyletic:
The
Distribution results genetic analysis: geographic pattern of analysis of genetic data of
The position of
Together with western and southern Anatolia, Crete is a biodiversity hotspot for
The current study revealed that based on morphological traits
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
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
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
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
The above scenario may not be unique and could be expected in other sibling species of Cretan
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
The song of
Mean syllable duration in recorded species from Crete (incl. Gavdos and Andikithira). See Suppl. material
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
Examples of syntopic occurrences of
Trap no. | Coll. date | Location | GPS Coordinates | Alt. (m) | Species 1 | Species 2 |
---|---|---|---|---|---|---|
FC111 | 30/10/1996 | Limni Kourna |
|
25 |
|
|
FC495 | 10/07/1997 | Limni Kourna |
|
25 |
|
|
FC70 | 20/08/1996 | Limni Kourna |
|
25 |
|
|
FC1536 | 05/08/2000 | Dikti Mt. |
|
1715 |
|
|
FC1606 | 02/10/2000 | Dikti Mt. |
|
1715 |
|
|
FC1655 | 09/01/2001 | Dikti Mt. |
|
1715 |
|
|
FC1602 | 15/09/2000 | Idi Mt. |
|
1910 |
|
|
FC1651 | 30/10/2000 | Idi Mt. |
|
1910 |
|
|
FC1916 | 12/06/2001 | Idi Mt. |
|
1910 |
|
|
FC17807 | 19/10/2018 | Lefka Mt. |
|
1200 |
|
|
Findings presented here, collected over the past 30 years, show that the known distribution range for five
Of the 14 species of
Pitfall traps are widely used to monitor ground dwelling invertebrates. For
Little information was available on the sympatric occurrences of different
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
Recently, a Red List for all European
Current IUCN Red List Status for
Species | Common name | IUCN RLA status* |
---|---|---|
|
Annamaria’s Marbled Bush-cricket |
|
|
Mount Ida Marbled Bush-cricket |
|
|
Cretan Marbled Bush-cricket |
|
|
Fer’s Marbled Bush-cricket |
|
|
Idi Marbled Bush-cricket |
|
Francis’s Marbled Bush-cricket | Not Evaluated | |
|
Skaronero Marbled Bush-cricket |
|
|
Giulia’s Marbled Bush-cricket |
|
|
Jacqueline’s Marbled Bush-cricket |
|
|
Hidden Marbled Bush-cricket |
|
|
Marianne’s Marbled Bush-cricket |
|
Marietherese’s Marbled Bush-cricket | Not Evaluated | |
|
Pale-legged Marbled Bush-cricket |
|
|
Smyrna Marbled Bush-cricket |
|
*:
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
Fourteen species of
This high concentration of endemic species of
Of the fourteen species of
Only one
Another interesting phylogeographical matter is that of
One more important question is, what caused the diversification of the 11 endemic
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 |
---|---|---|---|---|
|
|
2 | 3.3 Mya |
|
|
|
2 | 0.6 – 1.2 Mya |
|
|
|
2 | < 1 Mya |
|
|
|
4 | < 1 Mya | Fig. |
|
|
2 | 1.47 Mya | Vlachopoulos et al. (pers. Comm.) |
|
|
2 | 5.42 – 4.36 Mya |
*: but see also
East-west divergences are also evident in Cretan
Distribution pattern of Cretan
An example of another polyphyletic genus under investigation on Crete is
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
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
Dragan Chobanov would like to thank the grants supporting the
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
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
For helpful comments we are grateful to the reviewers Axel Hochkirch and Sigfrid Ingrisch.
List of localities, specimens, and repositories of
occurrence data (excel document)
The table contains all published and unpublished localities, specimens collected and their repositories for all
Examined specimens
specimen studied (excel document)
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
bioacoustic recordings (excel document)
table with detailed information on sound recordings and song data. The tab sound recordings summarises details of sound recordings of
Details of specimens used in phylogenetic analysis
table (excel document)
Details of specimens used for the Bayesian inference phylogenetic tree of 9
NADH2___ITIS_sequences
table (excel document)