Research Article |
Corresponding author: Kadir Boğaç Kunt ( chaetopelma@gmail.com ) Academic editor: Oana Teodora Moldovan
© 2019 Kadir Boğaç Kunt, Mert Elverici, Ersen Aydın Yağmur, Recep Sulhi Özkütük.
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:
Kunt KBB, Elverici M, Yağmur EA, Özkütük RS (2019) Kut gen. nov., a new troglomorphic spider genus from Turkey (Araneae, Dysderidae). Subterranean Biology 32: 95-109. https://doi.org/10.3897/subtbiol.32.46534
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A new genus of troglomorphic Dysderidae is described, based mainly on the morphology of copulatory organs. The new genus Kut gen. nov., with the type species Harpactocrates troglophilus Brignoli, 1978, also includes two recently discovered new species from the coastal Mediterranean Turkey: Kut izmiricus sp. nov. and K. dimensis sp. nov. All three species display troglomorphic traits, most distinct in K. dimensis sp. nov. Another genus-level trait is a characteristic simple type of bulb in males. Female copulatory organ shows similarity to the endemic Caucasian genus Cryptoparachtes Dunin, 1992 in paraspermatheca structure, whereas the male copulatory organ is unique among all known Dysderidae.
Anatolia, cave-dwelling, endemic, eyeless spider, MSS, troglobite
Dysderinae is one of the three subfamilies of the spider family Dysderidae, along with Harpacteinae and Rhodinae. Its members can be clearly distinguished by the unique morphology of sternum and labium joints, as well as by having claw tufts of setae on all tarsi and scopulae on posterior metatarsi (
Subfamily Dysderinae is quite diverse and includes the following genera (number of species given in square brackets): Cryptoparachtes Dunin, 1992 [3 spp.]; Dysdera Latreille, 1804 [285 spp.]; Dysderella Dunin, 1992 [2 spp.]; Dysderocrates Deeleman-Reinhold & Deeleman, 1988 [8 spp.]; Harpactocrates Simon, 1914 [13 spp.], Hygrocrates Deeleman-Reinhold, 1988 [5 spp.], Parachtes Alicata, 1964 [13 spp.], Stalitochara Simon, 1913 [1 sp.] and Tedia Simon, 1882 [2 spp.] (
Most of the genera of subfamily Dysderinae are endemic to the West Palearctic, with limited ranges. Cryptoparachtes and Dysderella are Caucasian endemics; Hygrocrates has a broader range from the Mediterranean Turkey and Caucasus; Harpactocrates and Parachtes are West European; Dysderocrates is East Mediterranean; and Tedia is known from Syria and Israel (
Many species of Dysderidae are adapted to life in the darkness and display troglomorphic traits (
Various authors doubted the generic placement of H. troglophilus, yet there have not been any attempts for a revision. In his original description,
This paper aims to contribute to the knowledge of the Dysderinae of Turkey by establishing a new genus based on troglomorphic Harpactocrates troglophilus and two other, new species recently discovered in Mediterranean Turkey.
Specimens were collected in different provinces of Turkey, either by hand collecting, pitfall, or MSS (Mesovoid Shallow Substratum) trapping and preserved in 70% ethanol. MSS trap design and application based on
Digital images of the left palp and vulva were taken with a Leica DFC295 digital camera attached to a Leica S8AP0 stereomicroscope. Five to 30 photographs were taken in different focal planes and combined using COMBINE ZP image stacking software. Photographic images were edited using ADOBE PHOTOSHOP CS6 and CORELDRAW HOME & STUDENT SUITE X7 was used to create the plates. All measurements are in mm. Terminology for the body parts and copulatory organ structures follows
Abbreviations: AL, abdominal length; CL, carapace length; CWmax, maximum carapace width; CWmin, minimum carapace width; AME, anterior median eyes; PLE, posterior lateral eyes; PME, posterior median eyes; AMEd, diameter of anterior median eyes; PLEd, diameter of posterior lateral eyes; PMEd, diameter of posterior median eyes; ChF, length of cheliceral fang; ChG, length of cheliceral groove; ChL, total length of chelicera (lateral external view); E embolus; T tegulum; Ms midspermatheca; Ps paraspermatheca; Ta, tarsus; Me, metatarsus, Ti, tibia; Pa, patella; Fe, femur; Tr, trochanter; C, coxa; D, dorsal; Pl, prolateral; Rl, retrolateral; V, ventral.
Depositories: AZMM, Zoology Museum of Alaşehir Vocational School, Celal Bayar University, Manisa, Turkey; ETZM, Eskişehir Technical University Zoology Museum, Eskişehir, Turkey.
Diagnosis. Sternum edge at labium-sternum joint approximately 2.5–3 times longer than the edges at gnathocoxa-sternum joints. All tarsi with claw tufts.
1 | Male | 2 |
– | Female | 8 |
2 | Distal extensions of tegulum simple | 3 |
– | Distal extensions of tegulum complex | 4 |
3 | Tegulum globular. Embolus long, sinuous. Conductor absent | Harpactocrates Simon, 1914 |
– | Tegulum pear-shaped. Embolus is a simple extension of tegulum |
Kut gen. nov. (Figs |
4 | Distal extensions divided with haematodocha | Dysdera Latreille, 1804 |
– | Distal extensions without haematodocha division | 5 |
5 | Tegulum Z-shaped, with two apophyses | Cryptoparachtes Dunin, 1992 |
– | Tegulum pear-shaped, straight or cylindrical | 6 |
6 | Tegulum pear-shaped, with two apophyses in addition to hook-shaped embolus | Hygrocrates Deeleman-Reinhold, 1988 |
– | Tegulum cylindrical | 7 |
7 | Tip of tegulum with a broad, well-developed, chitinized apophysis | Dysderocrates Deeleman-Reinhold & Deeleman, 1988 |
– | Tip of tegulum with a chitinized but underdeveloped, small apophysis | Parachtes Alicata, 1964 |
8 | Anterior spermatheca club-shaped | 9 |
– | Anterior spermatheca not club-shaped | 10 |
9 | Dorsal arc well-developed. Posterior diverticulum distinct |
Kut gen. nov. (Figs |
– | Posterior diverticulum indistinct | Cryptoparachtes Dunin, 1992 |
10 | Spermatheca with two distinct sections, distal and proximal | Hygrocrates Deeleman-Reinhold, 1988 |
– | Spermatheca without two distinct sections | 11 |
11 | Anterior diverticulum arc- or T-shaped | 12 |
– | Anterior diverticulum simple | Harpactocrates Simon, 1914 |
12 | Anterior diverticulum arc-shaped | 13 |
– | Anterior diverticulum T-shaped | Parachtes Alicata, 1964 |
13 | Anterior femora without numerous spines | Dysdera Latreille, 1804 |
– | Anterior femora with numerous spines | Dysderocrates Deeleman-Reinhold & Deeleman, 1988 |
Harpactocrates troglophilus Brignoli, 1978.
“Kut” is a Turkish word traceable to the old Turkic language with multiple meanings, such as “fortune (good or bad)”, “lifelong energy” or “vigour”, and more, depending on a dialect or a historical period. Gender: masculine.
The new genus resembles genera Dysderocrates, Harpactocrates and Hygrocrates in the general morphology of the male palp; however, it can be identified by the following characters:
In male copulatory organ of Kut gen. nov., tegulum is pear-shaped and embolus is a simple extension of a tegulum. Embolus represents the only distal extension of tegulum, any other apophyses are absent, unlike in Dysderocrates and Hygrocrates.
In Kut gen. nov., female copulatory organ is unique among the majority of Dysderinae genera in having the anteriorly located, club-shaped spermathecae, bearing a resemblance to those in Cryptoparachtes. Kut gen. nov. differs from Cryptoparachtes by having a well-developed dorsal arc and a distinct posterior diverticulum, as well as by the male copulatory organ (see
Anterior side of sternum (the side touching labium) is much more convex compared to other Dysderinae genera.
Large-sized Dysderinae spiders. Somatic characters as in other Dysderidae. Detailed description see under Kut troglophilus (Brignoli, 1978) comb. nov.
Harpactocrates troglophilus; Brignoli, 1978: 38, f. 4–5 (D♂).
H. troglophilus;
H. troglophilus;
11 ♂♂, 9 ♀♀ (ETZM and AZMM), Turkey: Konya Province, Beyşehir District, Kurucaova Town (37°40.483'N, 31°22.700'E), pitfall trap, August–October 2012, leg. E.A. Yağmur & O. Tutar; 1 ♂, Konya Province (ETZM), Beyşehir District, Kurucaova Town (37°40.450'N, 31°22.633'E), MSS trap, 08 August–10 October 2018, leg. E.A. Yağmur.
The simple structure of male palp in Kut troglophilus (Brignoli, 1978) is similar to that in some species of Harpactocrates, Harpactea, Parachtes, and Stalagtia. In K. troglophilus palp, the bulb gradually gets thinner and eventually gives rise to embolus, with a transition between tegulum and embolus rather indistinct, unlike in the other genera. Another unique feature of this genus is that the tip of the embolus is continuously bent posteriorly. In females, K. troglophilus vulva resembles that of the Caucasian genus Cryptoparachtes; however, in all three species of the latter (C. adzharicus, C. charitonowi and C. fedotovi), spermathecae are club-shaped. Paraspermatheca is distally spherical in all species of Cryptoparachtes, whereas in K. troglophilus paraspermathecae club shaped and more or less of equal length like the midspermatheca, unlike in three Cryptoparachtes species mentioned.
(♂/♀). AL 3.75–4.70/5.00–5.50; CL 3.50–4.48/3.50–4.00; CW 2.70–3.44/2.70–3.15; AMEd 0.14–0.18/0.12–0.16; PLEd 0.08–0.12/0.07–0.09; PMEd 0.08–0.11/0.07–0.12; ChF 0.95–1.25/0.96–1.05; ChG 0.55–0.80/0.60–0.70; ChL 1.60–2.10/1.60–1.75.
Carapace
hexagonal, reddish brown, with a smooth surface. Cephalic region darker than the thoracic region (Fig.
Labium and gnathocoxae
brown. Anterior sides of gnathocoxae chitinized at borders and dark brown. Labium anteriorly with a V-shaped incision. Sternum yellowish brown. Anterior side of sternum (touching labium) slightly convex (Fig.
Legs
yellowish, light brown. Leg coxa, trochanter, and femora darker in colour at joints. Femora and tibiae I and II prolaterally with spines; number of spines variable among individual spiders. Tibiae I and II with ventral spines. See Tables
Palp
(Fig.
Vulva
(Fig.
Legs | I | II | III | IV |
C | 1.98/1.84 | 1.80/1.55 | 1.20/1.00 | 1.25/1.15 |
Tr | 0.45/0.60 | 0.50/0.40 | 0.50/0.40 | 0.48/0.50 |
Fe | 3.60/3.20 | 3.35/2.90 | 2.80/2.50 | 3.50/3.35 |
Pa | 2.48/2.80 | 2.40/2.05 | 1.90/1.40 | 2.00/1.80 |
Ti | 3.04/2.30 | 2.95/2.35 | 2.35/2.00 | 3.05/2.35 |
Me | 3.04/1.85 | 3.00/2.25 | 2.75/2.55 | 3.50/3.35 |
Ta | 0.95/0.65 | 0.80/0.70 | 0.80/0.80 | 0.90/0.80 |
Total | 15.54/13.24 | 14.80/12.20 | 12.30/10.65 | 14.68/13.30 |
Legs | I | II | III | IV |
Fe | 4–6 Pl | 1, 1, 2–1, 2 Pl | 2–3 D | 3–5 Pl 3–5 D |
Pa | 0 | 0 | 1–2 Pl 1–3 Rl | 1 Pl 1–2 Rl |
Ti | 0–1 Pl 0–1 V | 0–1 Pl 1–2 V | 3–6 Pl 3–7 Rl 1, 1, 2 V | 3–6 Pl 3–6 Rl 1, 1, 2 V |
Me | 0 | 0 | 2–4 Pl 4–5 Rl 0–1, 1, 2 V | 3–5 Pl 4–7 Rl 1, 1, 2 V |
Holotype: 1 ♂ (ETZM), Turkey: İzmir Province, Kemalpaşa District, Vişneli Village, Nif Mountain (38°23.073'N, 27°21.614'E), MSS trap, 01 July–08 September 2015, leg. E.A. Yağmur, P. Mitov, S. Örgel & Ç. Altın. Paratypes: 1 ♀ (ETZM), 3 juveniles, 05 May–01 July 2015, leg. E.A. Yağmur, S. Örgel & S. Yaman, other data same as holotype; 3 subadults ♂♂, 07 September 2015–11 July 2016, leg. E.A. Yağmur, S. Örgel & S. Yaman, other data same as holotype; 1 ♂ (AZMM), 1 subadult ♀, 1 juvenile, 11 July–27 October 2016, leg. E.A. Yağmur, S. Örgel & S. Yaman, other data same as holotype; 1 ♂ (AZMM), 2 subadults ♀♀, 1 juvenile, 17 April–22 October 2017, leg. E.A. Yağmur, S. Örgel & S. Yaman, other data same as holotype; 3 subadults ♂♂ (AZMM), 09 June–20 October 2018, leg. E.A. Yağmur, S. Örgel & S. Yaman, other data same as holotype.
Male palp of Kut izmiricus sp. nov. resembles those of K. troglophilus and K. dimensis sp. nov.; in Kut izmiricus sp. nov. the bulb is more slender and exceeds embolus in length, unlike in the other two species. Female of Kut izmiricus sp. nov. differs from others by having a longer midspermatheca.
The specific name is a toponym that refers to the type locality, İzmir Province.
(Holotype ♂ / Paratype ♀). AL 6.56/5.80; CL 6.00/5.00; CW 4.32/3.60; AMEd 0.14/0.11; PLEd 0.08/0.10; PMEd 0.05/0.08; ChF 1.45/1.15; ChG 1.00/0.90; ChL 3.10/2.25.
Carapace
reddish brown, with thick dark brown contours at the borders. Fovea distinct, longitudinal (Fig.
Legs
brown. Coxae I, II and III darker in colour compared to other leg segments. See Tables
Palp
(Figs
Vulva
(Fig.
Legs | I | II | III | IV |
C | 2.80/2.15 | 2.56/2.00 | 1.50/1.35 | 1.70/1.25 |
Tr | 0.80/0.50 | 0.78/0.50 | 0.60/0.40 | 0.60/0.50 |
Fe | 5.84/4.25 | 5.00/3.75 | 4.40/3.25 | 5.44/4.20 |
Pa | 3.60/2.70 | 3.28/2.50 | 2.16/2.30 | 2.70/2.25 |
Ti | 4.80/3.25 | 4.64/3.50 | 3.50/3.75 | 4.20/3.75 |
Me | 4.40/3.00 | 4.32/3.25 | 4.30/3.50 | 5.70/4.25 |
Ta | 1.04/0.90 | 1.04/0.75 | 0.80/0.85 | 1.12/1.00 |
Total | 23.28/16.75 | 21.62/16.25 | 17.26/15.40 | 21.46/17.20 |
Holotype: 1 ♂ (ETZM), Turkey: Antalya Province, Alanya District, Kestel Town, Dim Cave (36°32.405'N, 32°6.603'E), hand collecting from stalactites and stalagmites, 04 January 2013, leg. K.B. Kunt & M. Elverici. Paratypes: 3 ♂♂, 5 ♀♀, 4 juveniles (ETZM), 1 ♂, 1 ♀ (AZMM), same data as holotype.
Male palp of Kut dimensis sp. nov. closely resembles that of K. troglophilus (Brignoli, 1978) comb. nov. Kut dimensis sp. nov. palp differs by having an embolus with tip sharply bent posteriorly, instead of gradually, and by having a more slender tegulum. Difference between Kut dimensis sp. nov. and K. izmiricus sp. nov. palps is more distinct, since in K. izmiricus sp. nov. tegulum/embolus ratio is higher compared to other species of the genus. K. dimensis sp. nov. females differ from other species by having longer paraspermathecae compared to midspermathecae, which is the opposite in the other two species.
The specific name is a toponym that refers to the type locality, Dim Cave located at Dim Valley in Alanya District.
(Holotype ♂ / Paratype ♀). AL 4.25/4.11; CL 4.40/4.25; CW 3.50/3.30; ChF 0.95/1.10; ChG 0.75/0.75; ChL 1.70/2.05.
Somatic traits of Kut dimensis sp. nov. resemble the other two species in general. Eye reduction at an extreme level, eyes almost absent (Fig.
Legs | I | II | III | IV |
C | 2.00/1.90 | 1.75/1.75 | 1.14/1.10 | 1.50/1.50 |
Tr | 0.55/0.35 | 0.35/0.40 | 0.30/0.45 | 0.50/0.50 |
Fe | 4.00/4.00 | 4.30/3.75 | 3.75/3.50 | 5.00/4.85 |
Pa | 2.75/2.50 | 2.80/2.50 | 1.75/1.75 | 2.40/2.30 |
Ti | 4.00/3.30 | 3.60/3.05 | 3.00/2.00 | 4.50/4.25 |
Me | 3.50/3.55 | 3.36/3.60 | 4.00/3.25 | 4.85/4.85 |
Ta | 0.75/0.85 | 0.50/0.80 | 0.70/0.75 | 0.90/0.65 |
Total | 17.55/16.45 | 16.66/15.85 | 14.64/12.80 | 19.65/18.90 |
Legs | I | II | III | IV |
Fe | 6–9 Pl 3–4 Rl | 5–7 Pl 0–4 Rl | 0–3 D 3–8 Pl 4–5 Rl | 3–4 D 4–6 Pl 3–4 Rl |
Pa | 0 | 0 | 1 Pl 3–4 Rl 0–1 V | 1 Pl 1 Rl 0–1 V |
Ti | 1, 1 V | 2–3 V | 6 Pl 6 Rl 1, 1, 2 V | 5–7 Pl 5–6 Rl 1, 1, 2 V |
Me | 1 V | 1 V | 3–5 Pl 5 Rl 1, 1, 2 V | 5–6 Pl 6 Rl 0–1, 1, 2 V |
Palp
(Fig.
Vulva
(Fig.
Kut dimensis sp. nov. is not the first troglomorphic species recorded from its type locality, the Dim Cave.
The Dim Valley, the valley where the Dim Cave located, has been studied by our team extensively in the recent past. There have been new species of Dysderidae discovered and described like Harpactea alanyana Özkütük et al., 2015 and Harpactea ballarini Kunt et al., 2013. Furthermore, the locality has been extensively sampled by the first author, as it was covered in his MSc thesis, titled “Harpacteinae Fauna of Antalya Province”. We provide these details to point out that Kut dimensis sp. nov. has not been found outside of the cave, in surface habitats, despite extensive sampling efforts, which is one of the reasons we consider this species a troglobiont.
With the description of Kut gen. nov., the number of genera belonging to spider family Dysderidae is now 25. Even though there are other genera known exclusively from hypogean habitats such as caves (for example Minotauria Kulczyński, 1903 and Folkia Kratochvil, 1970 from Harpacteinae; Stalita Schiödte, 1847 from Rhodinae), Kut gen. nov. is the first genus of Dysderidae ever discovered, in which all known species display troglomorphic traits such as reduced eyes or hypogean foraging. Kut troglophilus (Brignoli, 1978) is currently the only species of the Kut gen. nov. with recorded surface activity; however, it was also recorded from hypogean habitats: in MSS with this study and the Zindan Cave (
With this paper, we have resolved the discussion regarding the generic placement of Kut troglophilus. In our opinion, Kut gen. nov. has a unique and important position among Dysderidae due to its troglobitic and biogeographic affinities, coupled with the unique morphology of copulatory organs.
During our sampling efforts throughout the Mediterranean Turkey by using MSS traps, we have discovered further populations that might very likely belong to Kut gen. nov. as well. Moreover, we have collected subadult individuals from central parts of Anatolia with similar morphology, which may indicate a more or less continuous range of the genus throughout the Anatolia. We hope to reveal the specific identity of these populations, study their relationship to the known Aegean, Mediterranean, and Central Anatolian species, and the relationship between Kut gen. nov. and the Caucasian Cryptoparachtes in the near future. A high level of endemicity of Dysderinae around Anatolia at both species and genus levels indicates that many more taxa are still waiting to be discovered.
We are highly grateful to Dr. Miquel Arnedo (Barcelona, Spain) and Dr. Fulvio Gasparo (Trieste, Italy) for their valuable comments on the taxonomic position of the new genus, dating back to 2011. We thank Dr. Plamen Mitov (Sofia, Bulgaria), Dr. Semih Örgel (Manisa, Turkey), Serkan Yaman (Manisa, Turkey) and Çağatay Altın (Ardahan, Turkey) for their help during field trips. Dr. Victor Fet (West Virginia, USA) and Dr. Hayrettin İhsan Erkoç (Çanakkale, Turkey) kindly helped with nomenclatural acts. Mr. Savaş Kunt (Antalya, Turkey) provided accommodation during field studies in Alanya District. Special thanks go to Dr. Christo Deltshev (Sofia, Bulgaria) and Dr. Martina Pavlek (Barcelona, Spain) for reviewing the manuscript. The English of the final draft was kindly checked by Dr. Victor Fet (West Virginia, USA).