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Article

Contribution to the Knowledge on Distribution of Tardigrada in Turkey

by
Duygu Berdi
* and
Ahmet Altındağ
Department of Biology, Faculty of Science, Ankara University, 06100 Ankara, Turkey
*
Author to whom correspondence should be addressed.
Diversity 2020, 12(3), 95; https://doi.org/10.3390/d12030095
Submission received: 28 December 2019 / Revised: 29 February 2020 / Accepted: 4 March 2020 / Published: 6 March 2020
(This article belongs to the Special Issue Tardigrades Taxonomy, Biology and Ecology)
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Abstract

:
Tardigrades have been occasionally studied in Turkey since 1973. However, species number and distribution remain poorly known. In this study, distribution of Tardigrades in the province of Karabük, which is located in northern coast (West Black Sea Region) of Turkey, was carried out. Two moss samples were collected from the entrance of the Bulak (Mencilis) Cave. A total of 30 specimens and 14 eggs were extracted. Among the specimens; Echiniscus granulatus (Doyère, 1840) and Diaforobiotus islandicus islandicus (Richters, 1904) are new records for Karabük. Furthermore, this study also provides a current checklist of tardigrade species reported from Turkey, indicating their localities, geographic distribution and taxonomical comments.

1. Introduction

Caves are not only one of the most important forms of karst, but also one of the most unique forms of karst topography in terms of both size and formation characteristics, which are formed by mechanical melting and partly chemical erosion of water [1]. Most of the caves in Turkey were developed within the Cretaceous and Tertiary limestone, metamorphic limestone [2], and up to now ca. 40 000 karst caves have been recorded in Turkey. Although, most of these caves are found in the karstic plateaus zone in the Toros System, important caves, such as Kızılelma, Sofular, Gökgöl and Mencilis, have also formed in the Western Black Sea [3].
Some invertebrate communities can adapt well to these interesting underground formations, whose habitats are found rarely on earth [4,5]. Tardigrades, also known as water bears, are microscopic invertebrates that are one of these invertebrates that can survive in this environment with its harsh conditions. In general, tardigrades inhabit marine, freshwater and terrestrial environments (deserts, soil and leaf litters, mosses, lichens and liverworts, underground and marine caves throughout the world [6,7]. Until now, 30 families, 142 genera and nearly 1300 tardigrade species and 34 subspecies have been reported worldwide [8].
The tardigrade fauna of Turkey remains poorly known [9], with only few papers published since 1973. The most comprehensive studies were carried out by Maucci [10,11,12,13,14,15,16], Morgan [17], Binda [18], Kaczmarek et al. [9], Kharkevych and Sergeeva [19] and Ürkmez et al. [20]. To date 54 species and one subspecies have been reported from Turkey. Among them, eleven species and one subspecies belong to the genus Echiniscus C.A.S Shultze, 1840 [21] whereas only one species belongs to the genus Diaforobiotus Guidetti, Rebecchi, Bertolani, Jönsson, Kristensen and Cesari, 2016 [22].
This paper reports two tardigrade species, which were discovered for the first time from the Bulak (Mencilis) Cave, in Karabük. In addition, we also provide a full checklist of tardigrade species reported from Turkey with their localities, geographic distributions and taxonomical comments.

2. Materials and Methods

In March 2017, two moss samples were collected from the entrance of Bulak (Mencilis) Cave (41°16′30.1836″ N; 32°37′28.4628″ E), which is located nearly 8.5 km northwest of Safranbolu, the largest and most developed province of Karabük [23]. Bulak (Mencilis) Cave is the fourth largest cave in Turkey with a length of approximately 6 km [24,25].
One moss sample was collected from a rock which was found around the entrance of the Cave and the other from the wall of the entrance. Both samples were put into paper bags and let dry in the laboratory. They were later examined for terrestrial tardigrades using a standard method described by Dastych [26] with modifications described by Stec and Smolak [27]. The samples were placed in a beaker for rehydration with tap water in the laboratory. One day later after stirring, the water containing moss particles were poured into a 200 ml cylinder. After precipitation, approximately 50 ml of remaining sediment was taken into a glass petri dish. Tardigrades and eggs were placed from the sediment under the stereomicroscope (Leica MZ16 A). Tardigrades and their eggs were mounted on microscopic slides in Hoyer’s medium and observations and the photomicrographs were made using a phase-contrast microscope (Zeiss Axio Imager M1). The species were identified using the monograph published by Ramazotti and Maucci [7] and according to Guidetti et al. [22]. For genus abbreviations we referred to Perry et al. [28]. All slides, which were secured with a transparent nail polish, were stored in the Ankara University, Department of Biology, Aquatic Animals Research Laboratory.

3. Results

In total, 30 specimens and 14 eggs were found. Within those specimens, 10 belong to the genus Paramacrobiotus Guidetti, Schill, Bertolani, Dandekar and Wolf, 2009 [29] and they have not been identified to species level due to the absence of eggs. The remaining 20 specimens and 14 eggs were identified as Ech. granulatus (Doyère, 1840) [30] and Dia. i. islandicus (Richters, 1904) [31], which are new records for the province of Karabük.

Taxonomic Accounts

  • Phylum: Tardigrada Spallanzani, 1777 [32]
  • Class: Heterotardigrada Marcus, 1927 [33]
  • Order: Echiniscoidea Richters, 1926 [34]
  • Family: Echiniscidae Thulin, 1928 [35]
  • Genus: Echiniscus C.A.S. Schultze, 1840 [21]
  • Echiniscus granulatus
    • (Doyère, 1840) [30]
Material examined: Seven specimens.
Remarks: It has dorsal sculpture of polygonal shape (Figure 1C) that is composed of granules. Lateral appendices cirri A, C and D present (Figure 1A,B), which are in the form of long filaments, lateral cirri B absent. Cirri E in shape of spines. The dorsal appendices Cd and Dd as long spines. Fourth pair of legs with dentate collar (Figure 1D). All characteristics properties of this species correspond with the description in Ramazotti and Maucci [7]. The species is known from many localities in Europe, Turkey, North America, Africa and Asia [36].
Type locality: France [7]
  • Class: Eutardigrada Richters, 1926 [34]
  • Order: Macrobiotoidea Guil, Jørgensen and Kristensen, 2019 [37]
  • Family: Richtersiidae Guidetti, Rebecchi, Bertolani, Jönsson, Kristensen and Cesari, 2016 [22]
  • Genus: Diaforobiotus Guidetti, Rebecchi, Bertolani, Jönsson, Kristensen and Cesari, 2016 [22]
  • Diaforobiotus islandicus islandicus
    • (Richters, 1904) [31]
Material examined: In total 13 specimens and 14 eggs.
Remarks: Their body length is varied from 550 to 665 µm (Figure 2A). Buccal tube is thin and pharynx oval with two macroplacoids (Figure 2B); the first is longer than the second. Lunulas present and dentate (Figure 2C). The projections of the eggs are thin cones, almost spine-like (Figure 2D). Measurements of selected morphological characters for the smallest and largest examined specimens are given in Table 1. All characteristics properties of the species correspond with the description in Guidetti et al. [22]. Diaforobiotus i. islandicus (Richters, 1904) [31] was reported in Italy, Switzerland, Crete, Carpathians, Sweden, Norway, Iceland, Faroe Islands, Spitzbergen Archipelago, Greenland, America and Turkey [7,36]
Type locality: Iceland [7]
With our study, we added Karabük to the provinces in Turkey in which tardigrades have been recorded and thus, increased the number from 31 to 32 (see Figure 3 and Table 2 for details).

4. Discussion

We found Dia. i. islandicus (Richters, 1904) [31] in the province of Karabük, which is located in the Black Sea Region. It had previously been reported from İzmir, located in the Aegean Region of Turkey. This species has been identified from many localities in Europe, Africa, Indonesia, and North America [36].
Generally associated with limestone and alkaline bedrock, Ech. granulatus (Doyère, 1840) [30] has been reported from many localities in Europe, Africa, Caucasus, Georgia, Asia, Indonesia and North America [10,11,12,14,36,79]. Before our study, it had previously been reported from the provinces of Burdur, Bolu, Çorum, Erzurum, Kastamonu, Sivas, and Van in Turkey [36]. In the Poland, the species has been reported as a mountainous and sub-mountainous species and recorded between 500 and 1000 m asl on alkaline bedrocks [52]. In Turkey, this species was found in Çeltikçi Beli, Ilgaz Mountain, Boğazkale, Yıldızeli and Horasan regions [12]. Mostly usually recorded from 1000 m asl or above [10], this species has also been reported from the Trestede Karst plateau, in Italy, at between 200 to 300 m asl [10].
Although taxonomic and biogeographic studies on caves harboring tardigrades have been conducted for many years, there is still poor knowledge of their patterns of diversity. Only a few published papers center upon investigation for cave tardigrades. In 1996, Astatumen trinacriae (Arcidiacono, 1962) [80] was collected from the entrance of a cave located in Menglun, China [81]. Doryphoribius smokiensis Bartels, Nelson, Kaczmarek and Michalczyk, 2007 [82], Dor. longistipes Bartels, Nelson, Kaczmarek and Michalczyk, 2008 [83], and Dor. minimus Bartels, Nelson, Kaczmarek and Michalczyk, 2008 [83] were reported from the caves in the Great Smoky Mountains National Park in North America. In 2008, Minibiotus formosus Zawierucha, Dziamięcki, Jakubowska, Michalczyk and Kaczmarek, 2014 [84] has been collected from Gutmana Cave in the Gauja National Park [84]. In addition, Bartels et al. [85] described Mac. martini Bartels, Pilato, Lisi and Nelson, 2009 [86] from Gregorys Cave, North America. Nevertheless, one could think that cave invertebrates are more suitable for analysis, as the number of species is very low in comparison with to above ground habitats [5].
Up to now, tardigrades have been reported from 32 of the 82 Turkish provinces (see Table 2 for details). Among them, ten species have been described as endemic to Turkey (i.e., Cor. ceratophorus (Maucci, 1973) [10], Ech. dikenli Maucci, 1973 [10], Ech. osellai Maucci, 1975 [11], Ech. trojanus Maucci, 1973 [10], Urs. duranteae (Maucci, 1978) [12], Iso. macrodactylus (Maucci, 1978) [12], Gre. sismicus (Maucci, 1978) [12], Mil. beasleyi Kaczmarek, Jakubowska and Michalczyk, 2012 [9], Ita. placophorum Maucci, 1973 [10] and Ten. hystricogenitus (Maucci, 1978) [12]. In total, 54 valid species and one subspecies have been reported from Turkey. This number is quite inadequate when compared with other countries of smaller area (e.g., Romania ca 128, Poland ca. 102 and Italy ca. 233) [9,86,87,88,89]. Therefore, more samples are required to increase knowledge about distribution of tardigrades in Turkey.

Author Contributions

Methodology, D.B.; software, D.B.; validation, D.B. and A.A.; formal analysis, D.B. and A.A.; investigation, D.B. and A.A.; resources, D.B.; data curation, D.B.; writing—original draft preparation, D.B.; writing—review and editing, D.B.; visualization, D.B. All authors have read and agreed to the published version of the manuscript.

Funding

This research received no external funding.

Acknowledgments

We are grateful to Łukasz Kaczmarek for valuable comments and English improvements to this manuscript. We also wish to thank the Institute of Biotechnology at Ankara University for their contribution by providing technical and logistic support during our studies and the fieldwork.

Conflicts of Interest

The funders had no role in the design of the study; in the collection, analyses, or interpretation of data; in the writing of the manuscript, or in the decision to publish the results.

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Figure 1. Echiniscus granulatus: (A) Habitus, (B) dorsal plates, (C) detail of the cuticular ornamentation, (D) claws of the fourth pair of legs; using phase contrast microscopy.
Figure 1. Echiniscus granulatus: (A) Habitus, (B) dorsal plates, (C) detail of the cuticular ornamentation, (D) claws of the fourth pair of legs; using phase contrast microscopy.
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Figure 2. Diaforobiotus islandicus islandicus: (A) Habitus, (B) bucco-pharyngeal apparatus (lateral view), (C) claws of legs IV with large dentate lunula, (D) egg shell (midsection); using phase contrast microscopy.
Figure 2. Diaforobiotus islandicus islandicus: (A) Habitus, (B) bucco-pharyngeal apparatus (lateral view), (C) claws of legs IV with large dentate lunula, (D) egg shell (midsection); using phase contrast microscopy.
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Figure 3. The map of Turkey including all tardigrade species were reported from 32 provinces (marked in dark) From left to right: 1—Kırklareli, 2—İstanbul, 3—Sakarya, 4—Bolu, 5—Karabük *, 6—Kastamonu, 7—Sinop, 8—Rize, 9—Çanakkale, 10—Balıkesir, 11—Bursa, 12—Ankara, 13—Çorum, 14—Sivas, 15—Erzincan, 16—Tunceli, 17—Erzurum, 18—Ağrı, 19—İzmir, 20—Uşak, 21—Afyonkarahisar, 22—Konya, 23—Diyarbakır, 24—Bitlis, 25—Siirt, 26—Van, 27—Denizli, 28—Burdur, 29—Antalya, 30—Mersin, 31—Gaziantep, 32—Hakkari. Kaczmarek et al. [9]’s map is revised by adding the new provinces according to new records. * New province, which was added to the map with present study.
Figure 3. The map of Turkey including all tardigrade species were reported from 32 provinces (marked in dark) From left to right: 1—Kırklareli, 2—İstanbul, 3—Sakarya, 4—Bolu, 5—Karabük *, 6—Kastamonu, 7—Sinop, 8—Rize, 9—Çanakkale, 10—Balıkesir, 11—Bursa, 12—Ankara, 13—Çorum, 14—Sivas, 15—Erzincan, 16—Tunceli, 17—Erzurum, 18—Ağrı, 19—İzmir, 20—Uşak, 21—Afyonkarahisar, 22—Konya, 23—Diyarbakır, 24—Bitlis, 25—Siirt, 26—Van, 27—Denizli, 28—Burdur, 29—Antalya, 30—Mersin, 31—Gaziantep, 32—Hakkari. Kaczmarek et al. [9]’s map is revised by adding the new provinces according to new records. * New province, which was added to the map with present study.
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Table
Table 1. Measurements of selected morphological characters of largest and smallest specimens of Dia. i. islandicus and eggs (all values in μm, pt ratios in brackets).
Table 1. Measurements of selected morphological characters of largest and smallest specimens of Dia. i. islandicus and eggs (all values in μm, pt ratios in brackets).
CharacterSmallest SpecimenLargest Specimen
Body550665
Buccal tube length
Buccal tube external width
Buccal tube internal width		56.0
4.2 [7.5]
2.0 [3.6]		62.0
5.2 [8.4]
2.3 [3.7]
Stylet support insertion point16.9 [30.2]21.8 [35.2]
Macroplacoid row13.4 [23.9]20.0 [32.3]
First macroplacoid5.8 [10.4]9.3 [15.0]
Second macroplacoid4.0 [7.1]6.7 [10.8]
External claw I9.0 [16.1]13.2 [21.3]
Internal claw I14.5 [25.9]18.7 [30.2]
External claw II9.9 [17.7]14.0 [22.6]
Internal claw II15.2 [27.1]22.3 [36.0]
External claw III10.7 [19.1]14.3 [23.1]
Internal claw III16.1 [28.8]23.0 [37.1]
External claw IV15.8 [28.2]18.1 [29.2]
Internal claw IV22.6 [40.4]25.7 [41.5]
Egg diameter without processes94103
Diameter with processes133140
Process height9.6–11.711.9–12.1
Table
Table 2. The final list of all reported tardigrade species (marine and terrestrial) from Turkey.
Table 2. The final list of all reported tardigrade species (marine and terrestrial) from Turkey.
TaxaGeographic DistributionLocalities in Turkey
1. Acanthechiniscus
victor (Ehrenberg, 1853) [38]		Reported from many localities in Europe, North America and Greenland [36].This species was reported from Rize/Verçenik Mountain (2606 m asl) from Turkey [39]. And is described as an alpine taxon [7].
2. Bryodelphax parvulus
Thulin, 1928 [35]		Europe, Asia, Indonesia, Africa and N. America, S. America [36].It was recorded from Van, Turkey [12,14].
3. Cornechiniscus ceratophorus (Maucci, 1973) [10]Turkey [36].It is endemic to Turkey. It was reported in Burdur, Antalya [10,11,12,13].
4. Crn. cornutus (Richters, 1907) [40]Europe, Africa, Russia, Asia and N. America [36].Reported from Afyonkarahisar and Erzurum [11,13].
5. Crn. lobatus (Ramazzotti, 1943) [41]Europe, Russia, Asia and N. America [36].Reported from many localities in Çorum and Sivas [11,12].
6. Diaforobiotus islandicus islandicus (Richters, 1904) [31]Europe, Arctic, Africa, Asia, Indonesia and N. America [36].This species was reported from İzmir (Dikili) [10].
7. Dipodarctus subterraneus (Renaud-Debyser, 1959) [42]Faroe Islands, Malta, Bahamas, in Indian Ocean near Seychelles Maldivian Island [43,44], and Black Sea [19].This marine species has numerous reports from the Mediterranean Sea. In Turkey, it was found the Bosphorous at İstanbul in Black Sea [19].
8. Diphascon nobilei (Binda, 1969) [45]Europe, Russia, Australia and N. America [36].It was found in İzmir (Dikili) [12].
9. Doryphoribius flavus (Iharos, 1966) [46]Europe, Africa, Asia and Central America
[36].		Recorded at Ankara, Turkey [10].
10. Echiniscus blumi blumi Richters, 1903 [47] Reported from a few localities in Europe (Italy, Spain, Scotland, Switzerland, Turkey, Hungary, Greece) and N. America [36].It was reported from Ankara, Erzincan, and Van, Turkey [10,12,16]. As Ech. blumi blumi and as Ech. mediantus Marcus, 1930 [48] which is synonym of Ech. blumi blumi according to Gasiorek et al. [49].
11. Ech. canadensis Murray, 1910 [50]Europe, Africa, Asia, North and South America [36].It was reported from many localities from Turkey. Afyonkarahisar, Ağrı, Ankara, Bitlis, Çanakkale, Erzincan, İzmir, Kırklareli, Uşak, Van [10,11,12,16].
12. Ech. columinis Murray, 1911 [51]Europe and China [36].This palearctic species. It was only reported from Küçükkuyu/Çanakkale, Turkey [12].
13. Ech. dikenli Maucci, 1973 [10]Turkey [36].This endemic species was originally collected in Afyonkarahisar, Turkey [10,11,12].
14. Ech. granulatus (Doyère, 1840) [30]Europe, Africa, Asia, Indonesia and N. America [36].This species, reported from mountain and submountain regions [52] has been recorded from Bolu, Burdur, Çorum, Erzurum, Kastamonu, Sivas, Van [10,11,12,14], and now, in addition, from Karabük (this study).
15. Ech. merokensis merokensis Richters, 1904 [53]Europe, Africa, Asia, Russia, N. America and S. America [36].It was reported from İzmir, Turkey [10,12].
16. Ech. merokensis suecicus Thulin, 1911 [54]Reported from many localities in Europe, Africa and N. America [36].Reported from Bursa/Uludağ Mountain at 2200 m asl [10,12].
17. Ech. osellai Maucci, 1975 [11]Turkey [36].This endemic species has only reported from Antalya, Turkey [11,12].
18. Ech. testudo (Doyère, 1840) [30]Europe, Africa, Asia, N. America and S. America [9,12,36,55,56].It is cosmopolitan species. It was found in several localities in Afyonkarahisar, Ağrı, Antalya, Bitlis, Burdur, Çorum, Denizli, Diyarbakır, Erzincan, Erzurum, Gaziantep, Hakkari, Kastamonu, Mersin, Siirt, Sivas, Van and Konya, Turkey [10,11,17].
19. Ech. trisetosus Cuénot, 1932 [57]Europe, Africa, Asia and N. America [36].It was reported from Afyonkarahisar, Ağrı, Ankara, Balıkesir, Çanakkale, Çorum, Diyarbakır, Erzurum, Hakkari, İzmir, Kırklareli, Siirt, Uşak, Van, Turkey [10,11,12,16,17].
20. Ech. trojanus Maucci, 1973 [10]Turkey [36].This endemic species\was originally reported from Çanakkale, İzmir, Turkey [10,11,12].
21. Grevenius sismicus (Maucci, 1978) [12]Turkey and India [36].It was reported from Van, Turkey [12,14].
22. Hypsibius convergens (Urbanowicz, 1925) [58]Reported from many localities all over the world [36].This cosmopolitan species group requires the majority of records to be confirmed [9,59]. It has been reported from İzmir, Turkey [11,12].
23. Hys. microps Thulin, 1928 [35]Widespread throughout the world [36].This species is considered cosmopolitan, but a species group the majority of records need to be confirmed [9,59]. It has been reported from İzmir and Mersin, Turkey [11,12].
24. Isohypsibius macrodactylus (Maucci, 1978) [12]Italy, Turkey, Cyprus, Caucasus and Africa [36,60].A holarctic species was reported from Ağrı and Van, Turkey [9,12,14,36].
25. Iso. prosostomus Thulin, 1928 [35]Reported from many localities in Europe, Russia, Africa, Indonesia, N. America and S. America [36].Considered cosmopolitan, this species need to be re-described [9]. Reported from Ankara, Bolu, İzmir, Erzincan and Kastamonu, Turkey [10,11,12].
26. Itaquascon placophorum Maucci, 1973 [10]A few localities from Europe [36].It was originally described from Ankara and Sakarya, Turkey [10,11,12].
27. Macrobiotus crenulatus Richters, 1904 [53] Europe, Russia and N. America [36].Reported from Ankara. However, this species’ presence in Turkey needs to be confirmed [9,18,61].
28. Mac. echinogenitus Richters, 1904 [53]Europe, Russia, Arctic, Africa, Asia, Indonesia, Australia, New Zealand, N. America, Central America, S. America, Atlantic and Antarctica [36].Considered a cosmopolitan species group, it was reported from Ankara [12]. This species’ presence in Turkey needs to be verified [61].
29. Mac. hufelandi C.A.S. Schultze, 1834 [62]Europe, Russia, Africa, Asia, Indonesia, Australia, New Zealand, Pacific, N. America, Central America, S. America and Antarctica [36].A cosmopolitan species group that has been reported from Afyonkarahisar, Ankara, Antalya, Balıkesir, Bolu, Burdur, Denizli, İzmir, Kırklareli, Sakarya, Turkey [10,11].
30. Mac. pallarii Maucci, 1954 [63]Europe, Asia and N. America [36].Reported from three localities: Bolu, Kastamonu, Tunceli, Turkey [10,11,12].
31. Mac. persimilis Binda and Pilato, 1972 [64]Europe, Africa, Australia and N. America [36].Recorded from Balıkesir, Bolu, Diyarbakır, Erzincan, Sakarya, Uşak, Van, Turkey [12,14].
(Needs to be confirmed) [9].
32. Mac. spectabilis Thulin, 1928 [35]Europe, Turkey, Russia, N. America and S. America [36].Reported from the Verçenik Mountain/Rize, Turkey [39].
33. Megastygarctides sezginii Ürkmez, Ostrowska, Roszkowska, Gawlak, Zawierucha, Kristensen and
Kaczmarek, 2017 [20]		Black Sea in Turkey [20].Originally described from Sinop Bay, Turkey (Southern Black Sea) [20].
34. Mesobiotus harmsworthi (Murray, 1907) [65]Europe, Russia, Africa, Asia, Australia, Indonesia, New Zealand, N. America, Central America, S. America, Atlantic, Pacific and Antarctica [36].Cosmopolitan species, known from many localities throughout the world. It was found from Ağrı, Antalya, Bolu, Burdur, Bursa, Çorum, İzmir Kastamonu and Rize, Turkey [10,11,12,39].
35. Meb. nuragicus
(Pilato and Sperlinga, 1975)
[66]		Europe, Asia and Indonesia [36].Reported from Gaziantep, Turkey [12].
36. Meb. reinhardti (Michalczyk and Kaczmarek, 2003) [67]Cyprus and Turkey [9,67].Reported from Konya, Turkey [9].
37. Meb. stellaris (du Bois-Reymond Marcus, 1944) [68]Turkey, Greece and S. America (Brazil) [36].Recorded from Antalya, Turkey [11,12].
38. Milnesium beasleyi Kaczmarek, Jakubowska and Michalczyk, 2012 [9]Turkey [9].It is endemic species which was originally reported from Konya, Turkey [9].
39. Minibiotus intermedius (Plate, 1888) [69]Europe, Russia, Africa, Asia, Indonesia, Australia, New Zealand, Pacific, N. America, Central America, S. America, Indian and Antarctica [36].A cosmopolitan species group that has been reported from Ankara, Turkey [10,11,12].
40. Paramacrobiotus areolatus (Murray, 1907) [65]Europe, Africa, Asia, Indonesia, Australia, New Zealand, N. America, Central America and Atlantic [36].A cosmopolitan species group with an obscure their geographic distribution. It was reported from Uşak, Turkey [12].
41. Pam. richtersi (Murray, 1911) [51]Europe, Africa, Asia, Australia, New Zealand, N. America, Central America and S. America [36].A cosmopolitan species group reported from many localities in Turkey, including the provinces: Ankara, Antalya, Burdur, Çanakkale, Konya, Rize, İzmir, Van [9,10,11,14,39].
42. Parechiniscus chitonides Cuénot, 1926 [70]Europe, Russia and Africa [36].This palearctic species has been reported from Burdur and Antalya, Turkey [10,11,12].
43. Pilatobius patanei (Binda and Pilato, 1971) [71]Reported from Sicily and Turkey [36].This species was reported from an unidentified area in Anatolia [72].
44. Pseudechiniscus facettalis Petersen, 1951
[73]		Europe, Africa, New Zealand, N. America and S. America [36].Reported from İzmir, Turkey [10,11,12].
45. Pse. megacephalus Mihelčič, 1951 [74]Turkey and Austria [36].Only reported from Ankara, Turkey [10,11,12].
46. Pse. ramazottii ramazzottii Maucci, 1952 [75]Europe, Russia, North and South America [36].Recorded from the Verçenik Mountain/Rize, Turkey [39].
47. Pse. suillus (Ehrenberg, 1853) [38]Europe, Arctic, Africa, Asia, Indonesia, Indian, Australia, New Zealand, Pacific, N. America, Canada, Central America, S. America and Antarctica [36].Described as a cosmopolitan species group, this species was reported from Ankara and İzmir, Turkey [10].
48. Ramazzottius oberhaeuseri (Doyère, 1840) [30]Europe, Greenland, the Arctic and Antarctic, North and South America, Africa and New Zealand [36].A cosmopolitan species group that has been reported, from numerous locations in Turkey, including the provinces: Afyonkarahisar, Ağrı, Ankara, Antalya, Bolu, Çorum, Erzincan, Erzurum, Hakkari, Gaziantep, Kastamonu, Kırkareli, Uşak, Konya, Tunceli, Van and Rize [9,10,11,12,17,39]. However older records of this species need to be verified [9].
49. Richtersius coronifer (Richters, 1903) [47] Europe, Arctic, Africa, Asia, N. America and S. America [36].Reported from Ağrı, Çorum, İzmir and Van, Turkey [11,14].
50. Tanarctus ramazzottii Renaud-Mornant, 1975 [76]Gulf of Biscaglia, Coast of Britain, Mediterranean Sea [7,43] and Black Sea in Turkey [19].In Turkey, it has been reported from the Bosphorus of İstanbul in the Black Sea [19].
51. Tenuibiotus hystricogenitus (Maucci, 1978) [12]Turkey, Germany, Greece [36].It was recorded from Çorum and Erzurum, Turkey [12].
52. Ursulinius duranteae (Maucci, 1978) [12]Turkey [36].This endemic species was originally reported from Ağrı, Erzurum and Gaziantep, Turkey [12].
53. Urs. josephi (Iharos, 1964) [77]Europe [36].It was only recorded from Van, Turkey [12,14].
54. Urs. leithaicus (Iharos, 1966) [46]A few localities in Europe (Austria, Finland, Hungary) and Turkey [36].Reported from Van, Turkey [12].
55. Urs. torulosus (Mihelčič, 1959) [78]Turkey and Austria [36].Reported from İzmir, Turkey [11,12].

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Berdi, D.; Altındağ, A. Contribution to the Knowledge on Distribution of Tardigrada in Turkey. Diversity 2020, 12, 95. https://doi.org/10.3390/d12030095

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