Turkish Journal of Zoology Turk J Zool (2019) 43: 331-348 http://journals.tubitak.gov.tr/zoology/ © TÜBİTAK Research Article doi:10.3906/zoo-1902-5

Macrobiotus noemiae sp. nov., a new species (: hufelandi group) from Spain

1,2, 1 Milena ROSZKOWSKA *, Łukasz KACZMAREK  1 Department of Taxonomy and Ecology, Adam Mickiewicz University, Poznań, Poland 2 Department of Bioenergetics, Adam Mickiewicz University, Poznań, Poland

Received: 14.02.2019 Accepted/Published Online: 14.06.2019 Final Version: 01.07.2019

Abstract: A reexamination of the tardigrade collection from the Museo Nacional de Ciencias Naturales (MNCN) in Madrid was carried out. Specimens from the genera Milnesium, Macrobiotus, and Paramacrobiotus were verified in order to provide their correct identification and 15 taxa were identified. Moreover, a list of corrected species identifications from MNCN is provided. Fourteen specimens and 16 eggs, identified previously as Macrobiotus recens, were attributed to a species new for science. Macrobiotus noemiae sp. nov. is most similar to Mac. naskreckii and Mac. recens but it differs from them by details of egg morphology, mainly by the presence of thin flexible filaments on the tops of egg processes.

Key words: Europe, Eutardigrada, systematics, Tardigrada, taxonomy, water bears

1. Introduction not accepted by later authors. In 1993, Bertolani and The phylum Tardigrada consists of about 1300 species Rebecchi redescribed Mac. hufelandi and described (Guidetti and Bertolani, 2005; Degma and Guidetti, three new species of this complex, which resulted in 2007) that inhabit terrestrial, freshwater, and marine 17 species in the group in total. Later, many new taxa environments throughout the world (Ramazzotti and in the hufelandi group were described, and in the most Maucci, 1983 and later translation by Beasley, 1995; recent revision of the group, Kaczmarek and Michalczyk McInnes, 1994; Nelson et al., 2015).1 (2017) listed 47 species attributed to this complex. The tardigrade collection in the Museo Nacional Kaczmarek and Michalczyk (2017) defined the species de Ciencias Naturales (MNCN) in Madrid includes of the hufelandi group based on the presence of cuticular 3785 slides containing in total 19,099 specimens, eggs, pores, two macroplacoids (length sequence: 2 < 1) and a and exuviae. Samples of mosses, lichens, and leaf litter microplacoid in the pharynx, up to three bands of teeth containing and their eggs were collected in the oral cavity, Y-shaped claws, egg processes (single- between April 2001 and April 2003 by Noemi Guil López walled) in the shape of inverted chalices (although with and A. Sanchez Ruiz mainly from Spain, but a few samples some exceptions), and a smooth, porous, granulated, or were also collected from France and Belgium. reticulated egg shell. The genus Macrobiotus was described by C.A.S. In this paper, we reexamine tardigrades from the Schultze in 1834 as the first genus. At genera Macrobiotus; Mesobiotus Vecchi, Cesari, Bertolani, present it is the most species-rich genus in the phylum Jönsson, Rebecchi & Guidetti, 2016; Milnesium Doyère, Tardigrada.1 The term “Macrobiotus hufelandi group” was 1840; and Paramacrobiotus Guidetti, Schill, Bertolani, first used by Durante Pasa and Maucci (1979) for species Dandekar & Wolf, 2009 deposited on microscope slides resembling Macrobiotus hufelandi C.A.S Schultze, 1834. in the collection of the Museo Nacional de Ciencias Later, in 1990, Biserov partially revised the hufelandi Naturales, Madrid. Moreover, we provide a description group and proposed the subgenus Macrobiotus for this of a new Macrobiotus species belonging to the hufelandi group of species (Biserov, 1990a, 1990b), but it was group, previously identified as Mac. recens Cuénot, 1932. * Correspondence: [email protected] 1 Degma P, Bertolani R, Guidetti R (2009–2018). Actual checklist of Tardigrada species (v. 34: 30 June 2018), available online at www. tardigrada.modena.unimo.it/miscellanea/Actual%20checklist%20of%20Tardigrada.pdf. 331

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2. Materials and methods Order: Schuster, Nelson, Grigarick & 2.1. Microscopy and imaging Christenberry, 1980 According to information from the microscope slides, Superfamily: Macrobiotoidea Thulin, 1928 (in Marley specimens were mounted on microscope slides in Faure’s et al., 2011) medium and secured with a cover slip. Then slides were Family: Macrobiotidae Thulin, 1928 reexamined under an Olympus BX41 phase contrast light Genus: Macrobiotus C.A.S. Schultze, 1834 microscope (PCM) associated with a ARTCAM-300Mi Macrobiotus noemiae sp. nov., Tables 1 and 2, Figures digital camera (Olympus Corporation, Tokyo, Japan). 1–10 All figures were assembled in Corel Photo-Paint 2017. Macrobiotus recens Cuénot, 1932 (Guil, 2002), For deep structures that could not be fully focused in a probably also Macrobiotus recens (in Guil et al., 2009, Guil single photograph, a series of 2–10 images were taken and Sanchez-Moreno, 2013) every ~0.5 µm and then assembled into a single deep- Type material examined: 12 , 3 eggs (one with focus image manually in Corel Photo-Paint 2017. embryo) (holotype and 14 paratypes) on microscope slides in Faure’s medium. 2.2. Morphometrics and morphological nomenclature Additional material: 41°00′56.3″N, 3°39′0.6″W, 1040 All measurements are given in micrometers. Structures m a.s.l., Spain, 0.5 km from Gascones, road M-636, next were measured only if their orientation was suitable. to the pond, moss from rock, date 25 Oct. 2001 (2 eggs); Body length was measured from the anterior extremity 41°03′59.3″N, 3°31′9.6″W, 1160 m a.s.l., Spain, on the to the end of the body, excluding the hind legs. The Montejo city limit to La Hiruela, road M-137, leaf litter, types of buccopharyngeal apparatuses and claws were date 25 Oct. 2001 (1 specimen and 7 eggs); 40°50′22.1″N, classified according to Pilato and Binda (2010). All 3°57′16.8″W, 2020 m a.s.l., Spain, Parque Regional de measurements of adults and eggs of Macrobiotus were Peñalara, moss from rock, date 27 May 2001 (1 specimen prepared according to Kaczmarek and Michalczyk (2017). and 3 eggs); 41°06′34.3″N, 3°35′49.6″W, 1280 m a.s.l., Terminology describing the oral cavity armature (OCA) Spain, old road N-I, Dehesa Boyal, Robregordo, leaf litter, follows Michalczyk and Kaczmarek (2003), and OCA date 25 Oct. 2001 (1 egg). morphotypes are given according to Kaczmarek and Michalczyk (2017). The macroplacoid length sequence 3.1.1. Description of the new species was indicated according to Kaczmarek et al. (2014). The Animals (measurements and statistics in Table 1): pt ratio is the ratio of the length of a given structure to the Body transparent after fixation in Faure’s medium, eyes length of the buccal tube expressed as a percentage (Pilato, present in 33% of fixed specimens (Figure 1). Entire 1981). Genus abbreviations follow Perry et al. (2019). cuticle covered with conspicuous small, round pores Morphometric data were handled using the Parachela (0.8–1.1 µm in diameter) distributed randomly. Easily ver. 1.1 template available from the Tardigrada Register visible granulation present on legs I–IV (Figure 5). (Michalczyk and Kaczmarek, 2013). Tardigrade taxonomy Buccopharyngeal apparatus of the Macrobiotus type, follows Bertolani et al. (2014). with ventral lamina and ten peribuccal lamellae (Figures 1 and 2). Mouth anteroventral. Oral cavity armature of 2.3. Comparative material the patagonicus type, with only second and third bands For comparison with the new species, holotype and of teeth visible under light microscope (LM) (Figures 3a paratypes of Mac. naskreckii Bąkowski, Roszkowska, and 3b). The second band of teeth is composed of small Gawlak & Kaczmarek, 2016 from Łukasz Kaczmarek’s granules arranged in several rows, just above the third collection (Department of Animal Taxonomy and Ecology, band of teeth (Figures 3a and 3b, arrow). The third band of Faculty of Biology, Adam Mickiewicz University in Poznań) teeth is composed of three dorsal and three ventral teeth, were used. Moreover, in species identification as well as which appear under LM in the shape of transverse ridges revision of the slides from the MNCN collection, the keys (Figures 3a and 3b, indented arrowhead). Pharyngeal of Morek et al. (2016), Kaczmarek and Michalczyk (2017), bulb spherical with triangular apophyses, two rod- and Kaczmarek et al. (2017) and original descriptions/ shaped macroplacoids, and a triangular microplacoid. redescriptions were also used, i.e. Cuénot (1932), Maucci Macroplacoid length configuration 2 < 1 (Figure 2). The (1979), Ramazzotti and Maucci (1983), Michalczyk et first macroplacoid possesses a central constriction (Figure al. (2006, 2012a, 2012b), Kaczmarek et al. (2012), and 2, empty arrowhead). The second macroplacoid with Ciobanu et al. (2014, 2015). subterminal constriction (Figure 2, filled arrowhead). Claws of the hufelandi type, stout (Figures 4 and 6). 3. Results and discussion Primary branches with distinct accessory points. Lunules 3.1. Taxonomic account of the new species on claws I–III smooth, on claws IV dentate. Thin paired Phylum: Tardigrada Doyère, 1840 cuticular bars under claws I–III present (Figure 5, arrow). Class: Eutardigrada Richters, 1926 Other cuticular thickenings on legs absent.

332 ROSZKOWSKA and KACZMAREK / Turk J Zool

Table 1. Measurements (in µm) and pt values of selected morphological structures of individuals of Macrobiotus noemiae sp. nov. mounted in Faure’s medium (N – number of specimens/structures measured; Range – smallest and largest structures among all measured specimens; SD – standard deviation, pt –ratio of the length of a given structure to the length of the buccal tube expressed as a percentage).

Range Mean SD Holotype Character N µm pt µm pt µm pt µm pt Body length 12 248–625 – 434 955 106 – 625 – Buccopharyngeal tube Buccal tube length 12 34.2–56.4 – 44.8 – 6.3 – 56.4 – Stylet support insertion point 12 27.1–45.2 78.3–81.8 35.8 79.6 5.4 1.1 45.2 80.1 Buccal tube external width 12 4.8–9.6 14.0–17.5 7.1 15.8 1.3 1.0 9.6 17.0 Buccal tube internal width 9 3.1–7.0 9.1–14.3 5.2 11.5 1.1 1.3 6.4 11.3 Ventral lamina length 11 19.0–29.0 48.4–55.7 22.9 51.8 3.4 2.6 29.0 51.4 Placoid lengths Macroplacoid 1 12 6.5–14.1 19.0–25.0 10.3 22.7 2.0 1.5 14.1 25.0 Macroplacoid 2 12 5.5–10.6 15.2–20.1 7.5 16.5 1.6 1.5 10.4 18.4 Microplacoid 12 2.0–4.1 5.8–7.8 3.0 6.6 0.6 0.6 3.7 6.6 Macroplacoid row 12 13.4–25.9 39.2–47.1 19.1 42.4 3.7 2.2 25.9 45.9 Placoid row 12 16.1–30.7 47.1–57.9 23.1 51.2 4.5 3.1 30.7 54.4 Claw 1 lengths External primary branch 6 9.1–14.2 25.1–27.5 11.1 26.3 1.7 1.0 14.2 25.2 External secondary branch 6 8.7–12.7 22.5–26.9 10.5 24.8 1.4 1.8 12.7 22.5 Internal primary branch 7 7.8–13.7 22.0–26.2 10.5 24.0 1.8 1.6 ? ? Internal secondary branch 7 7.4–13.0 20.0–24.7 9.9 22.7 1.7 1.8 ? ? Claw 2 lengths External primary branch 8 9.5–17.2 27.8–32.6 14.0 30.3 2.4 1.9 16.0 28.4 External secondary branch 8 8.6–15.1 25.1–28.7 12.6 27.3 2.1 1.5 14.6 25.9 Internal primary branch 6 9.8–14.5 23.8–30.0 12.7 26.6 2.0 2.3 14.5 25.7 Internal secondary branch 6 8.9–14.2 21.7–29.1 12.0 25.2 1.9 2.7 13.1 23.2 Claw 3 lengths External primary branch 8 11.6–16.7 28.3–33.4 13.5 29.9 1.9 1.7 ? ? External secondary branch 8 10.0–15.0 23.8–30.0 11.9 26.3 1.9 1.9 ? ? Internal primary branch 5 8.1–11.6 23.7–28.0 10.1 25.2 1.3 1.7 ? ? Internal secondary branch 5 7.7–10.0 22.5–24.5 9.4 23.3 1.0 0.9 ? ? Claw 4 lengths Anterior primary branch 5 8.9–15.0 26.0–31.2 12.4 28.6 2.3 2.3 15.0 26.6 Anterior secondary branch 5 7.0–12.0 20.5–25.5 10.0 23.2 1.8 2.3 12.0 21.3 Posterior primary branch 4 9.7–16.0 25.7–30.9 12.8 28.3 2.6 2.1 16.0 28.4 Posterior secondary branch 4 7.5–12.9 21.0–24.9 10.3 22.7 2.2 1.7 12.9 22.9

Eggs (measurements and statistics in Table 2): Eggs contact (Figures 8 and 9). Processes in the shape of cones spherical, ornamented, and laid freely (Figures 7–10), with with blunt apex, but the apex sometimes divided on the top a chorion surface of the hufelandi type with reticulated (Figure 8, arrow). Terminal disc absent or not detectable surface, sometimes hardly visible (Figures 8 and 9). All under PCM. In the apical part of the processes few (2–5) processes situated close to one another, sometimes in long, thin, hair-like, and flexible filaments present (Figure

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Table 2. Measurements (in µm) of selected morphological structures of eggs of Macrobiotus noemiae sp. nov. mounted in Faure’s medium (N – number of specimens/structures measured, Range – smallest and largest structures among all measured eggs; SD – standard deviation).

Character N Range Mean SD Egg bare (without processes) diameter 3 100.6–105.7 103.6 2.7 Egg full (with processes) diameter 3 118.5–123.5 121.6 2.7 Process height 9 7.4–9.1 8.4 0.7 Process base width 9 6.8–8.1 7.5 0.5 Process base/height precentage 9 82%–94% 89% 5% Inter-process distance 9 1.5–2.2 1.9 0.3 Number of processes on the egg circumference 3 35–36 35.3 0.6

Figures 1–3a. Macrobiotus noemiae sp. nov.: 1- dorsoventral assembled image (holotype); 2- buccopharyngeal apparatus (dorsoventral assembled image), the empty arrowhead indicates the first macroplacoid with central constriction, the filled arrowhead indicates the second macroplacoid with a sub-terminal constriction (holotype); 3a- oral cavity armature, the arrow indicates teeth of the second band, the indented arrowhead indicates teeth of the third band (holotype). Scale bars in micrometers. All PCM.

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Figure 3b. Schematic illustrations of the oral cavity armature as seen under light microscopy (ventral and dorsal view, respectively).

Figures 4–6. Macrobiotus noemiae sp. nov.: 4- claws II with smooth lunules (holotype); 5- granulation on leg II (paratype); 6- claws IV with dentate lunules (holotype). Scale bars in micrometres. All PCM.

10, arrowheads). of an oral cavity armature of the patagonicus type Etymology: We dedicate this species to Noemi Guil (maculatus type in M. naskreckii); second macroplacoid López, a Spanish tardigradologist who collected the species with subterminal constriction; higher pt of stylet support and prepared the microscope slides that we examined. insertion point (78.3–81.8 in M. noemiae sp. nov. vs. Type locality: 40°51′29.1″N, 3°50′40.4″W, 1640 m 74.1–77.8 in M. naskreckii); lower pt of ventral lamina a.s.l., Spain, Puerto de la Morcuera, road M-611, leaf litter length (48.4–55.7 in M. noemiae sp. nov. vs. 57.7–63.1 in from meadow, date 16 May 2001, sample collector: Noemi M. naskreckii); egg chorion with visible reticulation under Guil López. LM (smooth in M. naskreckii); lack of terminal disc on the Type depositories: Holotype and 11 paratypes (slide: top of egg processes; presence of long, thin, hair-like, and 23.01/142), three eggs (slide: 23.01/2301), and additional flexible filaments on the top of processes; higher number of material are deposited at the Museo Nacional de Ciencias processes on the egg circumference (35–36 in M. noemiae Naturales in Madrid, Calle de José Gutiérrez Abascal, 2, sp. nov. vs. 27–32 in M. naskreckii); and larger egg bare 28006 Madrid, Spain. and full diameter (100.6–105.7 µm and 118.5–123.5 µm Differential diagnosis: Based on morphology of egg respectively in M. noemiae sp. nov. vs. 55.9–76.0 µm and processes, the new species is most similar to M. naskreckii 72.0–92.0 µm respectively in M. naskreckii). and M. recens, but differs specifically from: 2. Mac. recens, known from many localities, mainly 1. Mac. naskreckii, known only from its type locality in Europe (McInnes, 1994), by: egg chorion with visible in Mozambique (Bąkowski et al., 2016), by: presence reticulation under LM (slightly granulated in M. recens);

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Figures 7–10. Macrobiotus noemiae sp. nov. – eggs: 7- general view; 8–9- egg surface and processes, arrow indicates process with divided apical part; 10- long, hair-like, and flexible filaments present on the apical part of processes (arrowheads). Scale bars in micrometres. All PCM. presence of long, thin, hair-like, and flexible filaments on 3). Specimens identified previously asMac. recens were the top of processes; smaller egg full diameter (118.5– attributed to a species new for science described in this 123.5 µm in M. noemiae sp. nov. vs. 130.0–160.0 µm in M. paper. Examination of slides revealed this species also in recens); shorter processes (7.4–9.1 µm in M. noemiae sp. other samples, which may indicate a wider distribution of nov. vs. ca. 15.0 µm in M. recens). this species in Spain. The taxonomic status of Mac. recens 3.2. Remarks on the tardigrade collection from MNCN is still unclear and detailed redescription of this species is As part of the SYNTHESYS Project, 381 slides necessary (for more details see Pilato and Bertolani, 2004; containing species identified as Macrobiotus areolatus Stec et al., 2018). It was not possible to identify to the (now Paramacrobiotus areolatus (Murray, 1907)), Mac. species level most of the individuals attributed previously harmsworthi (now Mesobiotus harmsworthi (Murray, to Mac. harmsworthi and Mac. areolatus because in most 1907)), Mac. recens, Mac. richtersi (now Paramacrobiotus cases eggs were absent on examined slides. Only in one richtersi (Murray, 1911)), Milnesium eurystomum Maucci, case were eggs attributed to Mac. harmsworthi found to 1991, Mil. tardigradum Doyère, 1840, Mil. tetralamellatum belong to the newly described species – Mac. noemiae Pilato & Binda, 1991, and Milnesium sp. were verified in sp. nov. A similar situation was seen in the context of order to provide their correct identification (see Table specimens marked as the species Mac. richtersi, where in

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Table 3. Correct identifications of analyzed slides from the collection of the Museo Nacional de Ciencias Naturales in Madrid (Mac. – Macrobiotus; Mil. – Milnesium; Pam. – Paramacrobiotus).

Previous species No. of Correct species Slide number Coordinates (UTM) Additional information identification specimens identification 23.01/1019 Mil. tardigradum 30T0449865, 4551088 2 Mil. beasleyi 23.01/1035 Mil. tardigradum 30T0451167, 4542845 5 Mil. beasleyi 23.01/1039 Mil. tardigradum 30T0456366, 4546266 1 Mil. beasleyi 23.01/104 Mac. richtersi 30T0421842, 4507133 11 Pam. richtersi group 23.01/1045 Mil. tardigradum 30T0456046, 4534342 11 Mil. beasleyi 23.01/1047 Mil. tardigradum 30T0456046, 4534342 9 Mil. beasleyi 23.01/105 Mil. tardigradum 30T0421842, 4507133 1 Milnesium sp. Claws not clearly visible 23.01/1054 Mil. tardigradum 30T0452915, 4521442 3 Mil. beasleyi Specimens with Pyxidium 23.01/1064 Mil. tardigradum 30T0453780, 4514359 3 Mil. beasleyi tardigradum 23.01/1091 Mil. tardigradum 30T0408747, 4508333 19 Mil. beasleyi 23.01/475 Mil. tardigradum 30T0456046, 4534342 18 Mil. beasleyi 23.01/1432 Mil. tardigradum 30T0408747, 4508333 16 Mil. beasleyi 23.01/1435 Mil. tardigradum 30T0408747, 4508333 16 Mil. beasleyi 23.01/2146 Mil. tardigradum 30T0448826, 4525352 15 Mil. beasleyi 23.01/1429 Mil. tardigradum 30T0408747, 4508333 14 Mil. beasleyi 23.01/912 Mil. tardigradum 30T0449988, 4536504 13 Mi. beasleyi 23.01/1095 Mil. tardigradum 30T0408747, 4508333 11 Mil. beasleyi 23.01/2158 Mil. tardigradum 30T0408747, 4508333 11 Mil. beasleyi 23.01/395 Mil. tardigradum 30T0441624, 4533508 11 Mil. beasleyi 23.01/1437 Mil. tardigradum 30T0408747, 4508333 9 Mil. beasleyi 23.01/109 Mac. richtersi 30T0421842, 4507133 9 Pam. richtersi group 23.01/1073 Mil. tardigradum 30T0408747, 4508333 8 Mil. beasleyi 23.01/1085 Mil. tardigradum 30T0408747, 4508333 8 Mil. beasleyi 23.01/110 Mac. richtersi 30T0421842, 4507133 15 Pam. richtersi group 23.01/392 Mil. tardigradum 30T0441624, 4533508 8 Mil. beasleyi Mil. beasleyi, Mil. cf. Specimens with Pyxidium 23.01/497 Mil. tardigradum 30T0453780, 4514359 8 dornensis tardigradum 23.01/1406 Mil. tardigradum 30T0453780, 4514359 7 Mil. beasleyi 23.01/112 Mac. richtersi 30T0421842, 4507133 14 Pam. richtersi group 23.01/1155 Mac. harmsworthi 30T0414036, 4512512 1 Mesobiotus sp. 23.01/1483 Mil. tardigradum 30T0415046, 4507352 7 Mil. beasleyi 23.01/2118 Mil. tardigradum 30T0414036, 4512512 11 Mil.. cf. dornensis Rotifers and amoeba in gut 23.01/114 Mac. richtersi 30T0421842, 4507133 14 Pam. richtersi group Mil. cf. beatae, 23.01/2117 Mil. eurystomum 30T0414036, 4512512 9 Rotifers and amoeba in gut Milnesium sp. 23.01/477 Mil. tardigradum 30T0456046, 4534342 7 Mil. cf. dornensis 23.01/1139 Mil. eurystomum 30T0419895, 4519692 6 Mil. cf. beatae Rotifers in gut 23.01/115 Mac. richtersi 30T0421842, 4507133 15 Pam. richtersi group 23.01/1161 Mac. harmsworthi 30T0414036, 4512512 2 Mesobiotus sp. 23.01/1145 Mil. tardigradum 30T0419895, 4519692 6 Mil. cf. dornensis Rotifers in gut

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Table 3. (Continued).

23.01/2321 Mac. harmsworthi 30T0445333, 4540698 2 Mac. noemiae sp. nov. 23.01/323 Mac. harmsworthi 30T0432021, 4536661 1 Mesobiotus sp. 23.01/329 Mac. harmsworthi 30T0432021, 4536661 1 Mesobiotus sp. 23.01/116 Mac. richtersi 30T0421842, 4507133 14 Pam. richtersi group 23.01/429 Mac. harmsworthi 30T0445333, 4540698 2 Pam. richtersi group 23.01/434 Mac. harmsworthi 30T0448433, 4548409 2 Mesobiotus sp. 23.01/117 Mac. richtersi 30T0421842, 4507133 14 Pam. richtersi group 23.01/1372 Mil. tardigradum 30T0451428, 4554260 6 Mil. dornensis 23.01/440 Mac. harmsworthi 30T0449993, 4552368 4 Mesobiotus sp. 23.01/118 Mac. richtersi 30T0421842, 4507133 11 Pam. richtersi group 23.01/2134 Mil. tardigradum 30T0433525, 4510489 6 Mil. cf. dornensis 23.01/120 Mac. richtersi 30T0421842, 4507133 11 Pam. richtersi group 23.01/453 Mac. harmsworthi 30T0451264, 4553185 1 Mesobiotus sp. 23.01/5 Mac. harmsworthi 30T0405074, 4503305 2 Mesobiotus sp. 23.01/122 Mac. richtersi 30T0425612, 4504109 3 Pam. richtersi group 23.01/500 Mac. harmsworthi 30T0405074, 4503305 1 Mesobiotus sp. 23.01/551 Mac. harmsworthi 30T0409955, 4511632 1 Mesobiotus sp. 23.01/1362 Mil. tardigradum 30T0451428, 4554260 5 Mil. cf. dornensis 23.01/560 Mac. harmsworthi 30T0409955, 4511632 1 Mesobiotus sp. 23.01/605 Mac. harmsworthi 30T0419895, 4519692 2 Mesobiotus sp. 23.01/793 Mac. harmsworthi 30T0434152, 4519153 2 Mesobiotus sp. 23.01/8 Mac. harmsworthi 30T0405074, 4503305 4 Mesobiotus sp. 23.01/862 Mac. harmsworthi 30T0432021, 4536661 11 Mesobiotus sp. 23.01/1759 Mil. tardigradum 30T0445333, 4540698 7 Mil. beasleyi 23.01/865 Mac. harmsworthi 30T0432021, 4536661 4 Mesobiotus sp. 23.01/128 Mac. richtersi 30T0425612, 4504109 4 Pam. richtersi group 23.01/321 Mil. tardigradum 30T0439500, 4531983 7 Mil. beasleyi 23.01/13 Mac. richtersi 30T0408747, 4508333 11 Pam. richtersi group 23.01/130 Mac. richtersi 30T0425612, 4504109 1 Pam. richtersi group 23.01/894 Mil. tardigradum 30T0449988, 4536504 7 Mil. beasleyi 23.01/898 Mil. tardigradum 30T0449988, 4536504 7 Mil. beasleyi 23.01/1083 Mil. tardigradum 30T0408747, 4508333 6 Mil. beasleyi 23.01/1751 Mil. tardigradum 30T0445333, 4540698 6 Mil. beasleyi 23.01/135 Mac. richtersi 30T0425612, 4504109 5 Pam. richtersi group 23.01/1560 Mil. tardigradum 30T0428818, 4523346 3 Mil. cf. dornensis 23.01/1368 Mil. tardigradum 30T0451428, 4554260 1 Milnesium sp. 23.01/2124 Mil. tardigradum 30T0424219, 4509535 3 Mil. cf. dornensis 23.01/138 Mac. richtersi 30T0424921, 4511999 7 Pam. richtersi group 23.01/1933 Mil. tardigradum 30T0408747, 4508333 6 Mil. beasleyi 23.01/1937 Mil. tardigradum 30T0408747, 4508333 6 Mil. beasleyi 23.01/2073 Mil. tardigradum 30T0433525, 4510489 6 Mil. beasleyi 23.01/2162 Mil. tardigradum 30T0408747, 4508333 6 Mil. beasleyi Claws not clearly visible 23.01/1425 Mil. tardigradum 30T0408747, 4508333 2 Milnesium sp. Cuticle sculpture not visible

338 ROSZKOWSKA and KACZMAREK / Turk J Zool

Table 3. (Continued).

23.01/2179 Mil. tardigradum 30T0424663, 4507361 6 Mil. beasleyi Rotifers in gut 23.01/2225 Mil. tardigradum 30T0441624, 4533508 6 Mil. beasleyi 23.01/901 Mil. tardigradum 30T0449988, 4536504 6 Mil. beasleyi 23.01/143 Mil. tardigradum 30T0428818, 4523346 2 Milnesium sp. Identification not possible 23.01/1077 Mil. tardigradum 30T0408747, 4508333 5 Mil. beasleyi 23.01/1078 Mil. tardigradum 30T0408747, 4508333 5 Mil. beasleyi 23.01/1080 Mil. tardigradum 30T0408747, 4508333 5 Mil. beasleyi 23.01/144 Mac. richtersi 30T0424219, 4509535 8 Pam. richtersi group 23.01/1100 Mil. tardigradum 30T0408747, 4508333 5 Mil. beasleyi 23.01/1408 Mil. tardigradum 30T0453780, 4514359 5 Mil. beasleyi 23.01/1440 Mil. tardigradum 30T0408747, 4508333 5 Mil. beasleyi 23.01/2212 Mil. tardigradum 30T0447569, 4506108 5 Mil. beasleyi 23.01/149 Mac. richtersi 30T0428743, 4525608 1 Pam. richtersi group Larger number of specimens is 23.01/1521 Mil. tardigradum 30T0419517, 4521376 1 Milnesium sp. nov. needed to confirm 23.01/53 Mil. tardigradum 30T0415046, 4507352 3 Mil. dornensis Mil. beasleyi, Milnesium 23.01/999 Mil. tetralamellatum 30T0449462, 4550421 3 sp. 23.01/153 Mac. richtersi 30T0425098, 4527031 3 Pam. richtersi group Mil. beasleyi, Mil. cf. 23.01/479 Mil. tardigradum 30T0456046, 4534342 5 dornensis 23.01/904 Mil. tardigradum 30T0449988, 4536504 5 Mil. beasleyi 23.01/1079 Mil. tardigradum 30T0408747, 4508333 4 Mil. beasleyi 23.01/1082 Mil. tardigradum 30T0408747, 4508333 4 Mil. beasleyi Rotifers in gut 23.01/1143 Mil. tardigradum 30T0419895, 4519692 2 Mil. cf. dornensis 23.01/1572 Mil. tardigradum 30T0425098, 4527031 1 Milnesium sp. Probably a juvenile 23.01/158 Mac. richtersi 30T0425098, 4527031 1 Pam. richtersi group Exuviae, identification not 23.01/1584 Mil. tardigradum 30T0429715, 4531202 1 Milnesium sp. possible 23.01/1084 Mil. tardigradum 30T0408747, 4508333 4 Mil. beasleyi Exuviae, identification not 23.01/1615 Mil. tardigradum 30T0437885, 4512612 1 Milnesium sp. possible 23.01/1622 Mil. tardigradum 30T0437017, 4521195 1 Milnesium sp. Cuticle sculpture not visible 23.01/169 Mac. richtersi 30T0425098, 4527031 1 Pam. richtersi group 23.01/1692 Mil. tardigradum 30T0447569, 4506108 1 Milnesium sp. Buccal apparatus not visible 23.01/1703 Mil. tardigradum 30T0440726, 4512565 6 Mil. beasleyi 23.01/1335 Mil. tardigradum 30T0440726, 4512565 4 Mil. beasleyi 23.01/1338 Mil. tardigradum 30T0440726, 4512565 4 Mil. beasleyi 23.01/1443 Mil. tardigradum 30T0408747, 4508333 4 Mil. beasleyi 23.01/1988 Mil. tardigradum 30T0445333, 4540698 4 Mil. beasleyi 23.01/1992 Mil. tardigradum 30T0445333, 4540698 4 Mil. beasleyi 23.01/1074 Mil. tardigradum 30T0408747, 4508333 3 Mil. beasleyi 23.01/176 Mac. richtersi 30T0425098, 4527031 1 Pam. richtersi group 23.01/1249 Mil. tardigradum 30T043789, 4508465 3 Mil. beasleyi 23.01/1172 Mil. tardigradum 30T0419977, 4520916 2 Mil. cf. dornensis

339 ROSZKOWSKA and KACZMAREK / Turk J Zool

Table 3. (Continued).

23.01/123 Mil. tardigradum 30T0425612, 4504109 2 Mil. cf. dornensis 23.01/18 Mac. richtersi 30T0408747, 4508333 4 Pam. richtersi group 23.01/1810 Mil. tardigradum 30T0451428, 4554260 2 Mil. cf. dornensis 23.01/2183 Mil. tardigradum 30T0424219, 4509535 2 Mil. cf. dornensis 23.01/129 Mil. tardigradum 30T0425612, 4504109 3 Mil. beasleyi Specimens with Pyxidium 23.01/1534 Mil. tardigradum 30T0428818, 4523346 3 Mil. beasleyi tardigradum 23.01/1555 Mil. tardigradum 30T0428818, 4523346 3 Mil. beasleyi Male 23.01/1737 Mil. tardigradum 30T0441624, 4533508 3 Mil. beasleyi 23.01/1377 Mac. recens 30T0449865, 4551088 1 Mac. noemiae sp. nov. 23.01/1396 Mac. recens 30T0456366, 4546266 3 Mac. noemiae sp. nov. 23.01/142 Mac. recens 30T0428818, 4523346 11 Mac. noemiae sp. nov. 23.01/186 Mac. richtersi 30T0423927, 4521102 9 Pam. richtersi group 23.01/1824 Mac. recens 30T0456366, 4546266 1 Mac. noemiae sp. nov. 23.01/2295 Mac. recens 30T0419517, 4521376 3 Mac. noemiae sp. nov. 23.01/2301 Mac. recens 30T0428818, 4523346 3 Mac. noemiae sp. nov. 23.01/1749 Mil. tardigradum 30T0445333, 4540698 3 Mil. beasleyi 23.01/1874 Mil. tardigradum 30T0439500, 4531983 1 Mil. beasleyi 23.01/1871 Mil. tardigradum 30T0439500, 4531983 3 Mil. beasleyi 23.01/1886 Mil. tardigradum 30T0441624, 4533508 1 Milnesium sp Exuviae 23.01/2368 Mac. recens 30T0456366, 4546266 3 Mac. noemiae sp. nov. 23.01/2385 Mac. recens 30T0456366, 4546266 1 Mac. noemiae sp. nov. 23.01/91 Mac. recens 30T0419517, 4521376 1 Mac. noemiae sp. nov. 23.01/190 Mac. richtersi 30T0429715, 4531202 11 Pam. richtersi group 23.01/1955 Mil. tardigradum 30T0424663, 4507361 3 Mil. beasleyi Cuticle sculpture and claws not 23.01/1913 Mil. tardigradum 30T0439500, 4531983 1 Milnesium sp. visible 23.01/2019 Mil. tardigradum 30T0451167, 4542845 3 Mil. beasleyi 23.01/753 Mil. tardigradum 30T0429715, 4531202 3 Mil. beasleyi Specimens with Pyxidium 23.01/1925 Mil. tardigradum 30T0441624, 4533508 1 Milnesium sp. tardigradum 23.01/916 Mil. tardigradum 30T0449988, 4536504 3 Mil. beasleyi 23.01/1104 Mil. tardigradum 30T0408747, 4508333 2 Mil. beasleyi 23.01/1333 Mil. tardigradum 30T0440726, 4512565 2 Mil. beasleyi 23.01/1393 Mil. tardigradum 30T0451167, 4542845 2 Mil. beasleyi Cuticle sculpture and claws not 23.01/1978 Mil. tardigradum 30T0440726, 4512565 1 Milnesium sp. visible 23.01/1551 Mil. tardigradum 30T0428818, 4523346 2 Mil. beasleyi 23.01/1592 Mil. tardigradum 30T043789, 4508465 2 Mil. beasleyi 23.01/1733 Mil. tardigradum 30T0441624, 4533508 2 Mil. beasleyi 23.01/1735 Mil. tardigradum 30T0441624, 4533508 2 Mil. beasleyi 23.01/1830 Mil. tardigradum 30T0452915, 4521442 2 Mil. beasleyi Mil. beasleyi, Mi. cf. 23.01/586 Mil. tardigradum 30T0415046, 4507352 2 dornensis

340 ROSZKOWSKA and KACZMAREK / Turk J Zool

Table 3. (Continued).

23.01/2053 Mil. tardigradum 30T0419977, 4520916 1 Milnesium sp. Identification not possible 23.01/643 Mil. tardigradum 30T0424663, 4507361 2 Mil. cf. dornensis 23.01/1142 Mil. eurystomum 30T0419895, 4519692 1 Mil. cf. dornensis Rotifers in gut 23.01/1837 Mil. tardigradum 30T0453780, 4514359 2 Mil. beasleyi Rotifers in gut 23.01/1976 Mil. tardigradum 30T0440726, 4512565 2 Mil. beasleyi 23.01/1152 Mil. tardigradum 30T0414036, 4512512 1 Mil. cf. dornensis 23.01/1189 Mil. tardigradum 30T0428818, 4523346 1 Mil. cf. dornensis 23.01/1523 Mil. tardigradum 30T0424663, 4507361 1 Mil. cf. dornensis 23.01/2120 Mil. tardigradum 30T0419977, 4520916 1 Milnesium sp. Cuticle sculpture not visible Cuticle sculpture and claws not 23.01/2122 Mil. tardigradum 30T0424663, 4507361 2 Milnesium sp. visible 23.01/1525 Mil. tardigradum 30T0424921, 4511999 1 Mil. cf. dornensis 23.01/1795 Mil. tardigradum 30T0451428, 4554260 1 Mil. cf. dornensis 23.01/2154 Mil. tardigradum 30T0451167, 4542845 2 Mil. beasleyi 23.01/2213 Mil. tardigradum 30T0440726, 4512565 2 Mil. beasleyi 23.01/2239 Mil. tardigradum 30T0451167, 4542845 2 Mil. beasleyi 23.01/250 Mil. tardigradum 30T0437885, 4512612 2 Mil. beasleyi Specimens with Pyxidium 23.01/1799 Mil. tardigradum 30T0451428, 4554260 1 Mil. cf. dornensis tardigradum 23.01/343 Mil. tardigradum 30T0432021, 4536661 2 Mil. beasleyi 23.01/384 Mil. tardigradum 30T0448826, 4525352 2 Mil. beasleyi 23.01/623 Mil. tardigradum 30T0410875, 4511586 2 Mil. beasleyi 23.01/1805 Mil. tardigradum 30T0451428, 4554260 1 Mil. cf. dornensis 23.01/763 Mil. tardigradum 30T0430540, 4504476 2 Mil. beasleyi 23.01/2051 Mil. tardigradum 30T0419977, 4520916 1 Milnesium sp. Cuticle scalpture not visible 23.01/910 Mil. tardigradum 30T0449988, 4536504 2 Mil. beasleyi 23.01/2055 Mil. tardigradum 30T0419977, 4520916 1 Milnesium sp. Claws not visible 23.01/2209 Mil. tardigradum 30T0437017, 4521195 1 Mil. berladnicorum 23.01/1070 Mil. tardigradum 30T0455633, 4513774 1 Mil. beasleyi 23.01/1102 Mil. tardigradum 30T0408747, 4508333 1 Mil. beasleyi 23.01/2060 Mil. tardigradum 30T0424219, 4509535 1 Mil. cf. dornensis 23.01/1135 Mil. tardigradum 30T0415046, 4507352 1 Mil. beasleyi 23.01/2110 Mil. tardigradum 30T0409955, 4511632 1 Mil. cf. dornensis 23.01/2155 Mil. tardigradum 30T0452938, 4533022 1 Mil. cf. dornensis 23.01/1321 Mil. tardigradum 30T0447569, 4506108 1 Mil. beasleyi Male 23.01/2259 Milnesium sp. 1 Milnesium sp. Exuviae 23.01/228 Mac. richtersi 30T0429195, 4530946 1 Pam. richtersi group 23.01/237 Mac. richtersi 30T0427136, 4530446 3 Pam. sklodowskae 23.01/239 Mac. richtersi 30T0430540,4504476 2 Pam. richtersi group 23.01/243 Mac. richtersi 30T0433525, 4510489 1 Pam. richtersi group 23.01/246 Mac. richtersi 30T0433525, 4510489 4 Pam. richtersi group 23.01/1400 Mil. tardigradum 30T0456046, 4534342 1 Mil. beasleyi 23.01/1426 Mil. tardigradum 30T0408747, 4508333 1 Mil. beasleyi

341 ROSZKOWSKA and KACZMAREK / Turk J Zool

Table 3. (Continued).

23.01/1427 Mil. tardigradum 30T0408747, 4508333 1 Mil. beasleyi 23.01/1485 Mil. tardigradum 30T0419895, 4519692 1 Mil. beasleyi 23.01/290 Mac. richtersi 30T0434152, 4519153 3 Pam. cf. sklodowskae Egg damaged 23.01/293 Mac. richtersi 30T0437017, 4521195 13 Pam. richtersi group 23.01/295 Mac. richtersi 30T0437017, 4521195 5 Pam. richtersi group 23.01/298 Mac. richtersi 30T0437017, 4521195 7 Pam. richtersi group 23.01/318 Mac. richtersi 30T0439500, 4531983 11 Pam. richtersi group 23.01/320 Mac. richtersi 30T0439500, 4531983 1 Pam. richtersi group 23.01/1547 Mil. tardigradum 30T0428818, 4523346 1 Mil. beasleyi 23.01/1762 Mil. tardigradum 30T0445333, 4540698 1 Mil. beasleyi 23.01/1831 Mil. tardigradum 30T0455252, 4529638 1 Mil. beasleyi 23.01/1834 Mil. tardigradum 30T0453780, 4514359 1 Mil. beasleyi 23.01/344 Mac. richtersi 30T0435667, 4533269 1 Pam. sklodowskae 23.01/345 Mac. richtersi 30T0435667, 4533269 21 Pam. sklodowskae 23.01/1877 Mil. tardigradum 30T0439500, 4531983 1 Mil. beasleyi Cuticle sculpture not visible 23.01/349 Mac. richtersi 30T0435667, 4533269 3 Pam. sklodowskae 23.01/352 Mac. richtersi 30T0447569, 4506108 11 Pam. richtersi group 23.01/353 Mac. richtersi 30T0447569, 4506108 14 Pam. richtersi group 23.01/355 Mac. richtersi 30T0447569, 4506108 13 Pam. richtersi group 23.01/357 Mac. richtersi 30T0447569, 4506108 11 Pam. richtersi group 23.01/359 Mac. richtersi 30T0447569, 4506108 7 Pam. richtersi group 23.01/364 Mac. richtersi 30T0449010, 4519466 17 Pam. richtersi group 23.01/374 Mac. richtersi 30T0443836, 4515649 3 Pam. richtersi group 23.01/383 Mac. richtersi 30T0448826, 4525352 6 Pam. richtersi group 23.01/1901 Mil. tardigradum 30T0408747, 4508333 1 Mil. beasleyi 23.01/388 Mac. richtersi 30T0442434, 4524168 6 Pam. richtersi group 23.01/39 Mac. richtersi 30T0409889, 4510268 5 Pam. richtersi group No eggs 23.01/391 Mac. richtersi 30T0441624, 4533508 3 Pam. richtersi group 23.01/1917 Mil. tardigradum 30T0439500, 4531983 1 Mil. beasleyi 23.01/394 Mac. richtersi 30T0441624, 4533508 2 Pam. richtersi group Specimens with Pyxidium 23.01/1922 Mil. tardigradum 30T0441624, 4533508 1 Mil. beasleyi tardigradum 23.01/411 Mac. richtersi 30T0441091, 4537401 2 Pam. richtersi group 23.01/1985 Mil. tardigradum 30T0441624, 4533508 1 Mil. beasleyi 23.01/435 Mac. richtersi 30T0448433, 4548409 3 Pam. richtersi group 23.01/442 Mac. richtersi 30T0449462, 4550421 3 Pam. richtersi group 23.01/443 Mil. tardigradum 30T0449462, 4550421 1 Milnesium sp Claws not clearly visible 23.01/447 Mac. richtersi 30T0450100, 4552133 8 Pam. richtersi group 23.01/450 Mac. richtersi 30T0449865, 4551088 5 Pam. richtersi group Cuticle sculpture and claws not 23.01/451 Mil. tardigradum 30T0449865, 4551088 4 Milnesium sp. visible 23.01/454 Mac. richtersi 30T0451264, 4553185 1 Pam. richtersi group 23.01/456 Mac. richtersi 30T0452720, 4545827 1 Pam. richtersi group

342 ROSZKOWSKA and KACZMAREK / Turk J Zool

Table 3. (Continued).

23.01/457 Mac. richtersi 30T0451167, 4542845 16 Pam. richtersi group 23.01/2028 Mil. tardigradum 30T0451180, 4513830 1 Mil. beasleyi 23.01/46 Mac. richtersi 30T0415046, 4507352 8 Pam. richtersi group 23.01/462 Mac. richtersi 30T0451167, 4542845 9 Pam. richtersi group 23.01/466 Mac. richtersi 30T0456366, 4546266 11 Pam. richtersi group 23.01/2085 Mil. tardigradum 30T0440726, 4512565 1 Mil. beasleyi 23.01/2164 Mil. tardigradum 30T0409955, 4511632 1 Mil. cf. dornensis 23.01/2139 Mil. tardigradum 30T0439500, 4531983 1 Mil. beasleyi Cuticle sculpture and claws not 23.01/48 Mil. tardigradum 30T0415046, 4507352 1 Milnesium sp. visible 23.01/481 Mac. richtersi 30T0452915, 4521442 6 Pam. richtersi group 23.01/2152 Mil. tardigradum 30T0451264, 4553185 1 Mil. beasleyi 23.01/484 Mac. richtersi 30T0452915, 4521442 7 Pam. richtersi group 23.01/486 Mac. richtersi 30T0455252, 4529638 7 Pam. richtersi group 23.01/489 Mac. richtersi 30T0454613, 4524479 1 Pam. richtersi group 23.01/493 Mac. richtersi 30T0453780, 4514359 7 Pam. richtersi group 23.01/2157 Mil. tardigradum 30T0454613, 4524479 1 Mil. beasleyi 23.01/51 Mac. richtersi 30T0415046, 4507352 6 Pam. richtersi group 23.01/516 Mac. richtersi 30T0408747, 4508333 3 Pam. richtersi group 23.01/523 Mac. richtersi 30T0408747, 4508333 1 Pam. richtersi group 23.01/529 Mac. richtersi 30T0408747, 4508333 8 Pam. richtersi group 23.01/2175 Mil. tardigradum 30T0414036, 4512512 1 Mil. cf. dornensis 23.01/539 Mac. richtersi 30T0408621, 45072201 4 Pam. richtersi group 23.01/543 Mac. richtersi 30T0408621, 45072201 5 Pam. richtersi group 23.01/2221 Mil. tardigradum 30T0441624, 4533508 1 Mil. cf. dornensis 23.01/55 Mac. richtersi 30T0415046, 4507352 11 Pam. richtersi group 23.01/553 Mac. richtersi 30T0409955, 4511632 3 Pam. richtersi group 23.01/576 Mac. richtersi 30T0409889, 4510268 6 Pam. richtersi group 23.01/58 Mac. richtersi 30T0415046, 4507352 6 Pam. sklodowskae 23.01/580 Mac. richtersi 30T0409889, 4510268 5 Pam. richtersi group 23.01/2232 Mil. tardigradum 30T0445333, 4540698 1 Mil. cf. dornensis Cuticle sculpture not visible 23.01/2236 Mil. tardigradum 30T0449462, 4550421 1 Mil. cf. dornensis Male 23.01/591 Mac. richtersi 30T0414691, 4504440 17 Pam. richtersi group 23.01/592 Mil. tardigradum 30T0414691, 4504440 1 Milnesium sp. Cuticle sculpture not visible 23.01/593 Mac. richtersi 30T0414691, 4504440 13 Pam. richtersi group 23.01/547 Mil. tardigradum 30T0409955, 4511632 1 Mil. cf. dornensis 23.01/599 Mil. tardigradum 30T0419895, 4519692 1 Milnesium sp. Cuticle sculpture not visible 23.01/60 Mil. tardigradum 30T0415046, 4507352 1 Milnesium sp. Claws not clearly visible 23.01/61 Mac. richtersi 30T0415046, 4507352 4 Pam. richtersi group 23.01/613 Mil. tardigradum 30T0414036, 4512512 1 Milnesium sp. Claws not clearly visible 23.01/620 Mac. richtersi 30T0410875, 4511586 1 Pam. richtersi group 23.01/622 Mac. richtersi 30T0410875, 4511586 5 Pam. richtersi group 23.01/2167 Mil. tardigradum 30T0418705, 4507089 1 Mil. beasleyi

343 ROSZKOWSKA and KACZMAREK / Turk J Zool

Table 3. (Continued).

23.01/585 Mil. tardigradum 30T0409962, 4514249 1 Mil. cf. dornensis Male Adult specimens needed for final 23.01/628 Mac. richtersi 30T0419517, 4521376 1 Paramacrobiotus sp. identification 23.01/634 Mac. richtersi 30T0421842, 4507133 9 Pam. richtersi group 23.01/641 Mac. richtersi 30T0425612, 4504109 1 Pam. richtersi group 23.01/598 Mil. eurystomum 30T0419895, 4519692 1 Mil. cf. dornensis Male Final eggs identification not 23.01/644 Mac. richtersi 30T0424921, 4511999 4 Pam. cf. sklodowskae possible Final eggs identification not 23.01/646 Mac. richtersi 30T0424921, 4511999 11 Pam. cf. sklodowskae possible 23.01/648 Mac. richtersi 30T0424219, 4509535 6 Pam. richtersi group 23.01/657 Mac. richtersi 1 Pam. richtersi group 23.01/659 Mac. richtersi 1 Pam. richtersi group 23.01/66 Mac. richtersi 30T0414691, 4504440 4 Pam. richtersi group 23.01/661 Mac. richtersi 7 Pam. richtersi group 23.01/679 Mac. richtersi 30T0425098, 4527031 2 Pam. richtersi group 23.01/68 Mac. areolatus 30T0419895, 4519692 1 Pam. areolatus group 23.01/688 Mac. richtersi 30T0425098, 4527031 1 Pam. richtersi group 23.01/698 Mac. richtersi 30T0425098, 4527031 1 Pam. richtersi group 23.01/706 Mac. richtersi 30T0425098, 4527031 3 Pam. richtersi group 23.01/717 Mac. richtersi 30T0425098, 4527031 3 Pam. richtersi group 23.01/721 Mac. richtersi 30T0425098, 4527031 8 Pam. richtersi group 23.01/625 Mil. tardigradum 30T0419977, 4520916 1 Mil. dornensis Rotifers in gut 23.01/743 Mac. richtersi 30T0429715, 4531202 1 Pam. cf. sklodowskae 23.01/746 Mac. richtersi 30T0429715, 4531202 7 Pam. cf. sklodowskae 23.01/750 Mac. richtersi 30T0429715, 4531202 16 Pam. cf. sklodowskae 23.01/2461 Mil. tardigradum 1 Mil. beasleyi 23.01/752 Mac. richtersi 30T0429715, 4531202 14 Pam. richtersi group 23.01/2462 Mil. tardigradum 1 Mil. beasleyi 23.01/754 Mac. richtersi 30T0429715, 4531202 17 Pam. richtersi group 23.01/2474 Mil. tardigradum 1 Mil. beasleyi 23.01/758 Mac. richtersi 30T0435142, 4506143 4 Pam. cf. sklodowskae Eggs damaged 23.01/759 Mil. tardigradum 30T0435142, 4506143 2 Milnesium sp. Cuticle sculpture not visible 23.01/762 Mac. richtersi 30T0430540, 4504476 1 Pam. richtersi group 23.01/332 Mil. tardigradum 30T0432021, 4536661 1 Mil. beasleyi 23.01/767 Mil. tardigradum 30T0433525, 4510489 1 Milnesium sp. 23.01/779 Mac. richtersi 30T0437885, 4512612 5 Pam. sklodowskae 23.01/78 Mac. richtersi 30T0410875, 4511586 7 Pam. richtersi group 23.01/338 Mil. tardigradum 30T0432021, 4536661 1 Mil. beasleyi 23.01/787 Mac. richtersi 30T0434152, 4519153 1 Pam. richtersi group 23.01/794 Mac. richtersi 30T0434152, 4519153 1 Pam. richtersi group 23.01/800 Mac. richtersi 30T0434152, 4519153 1 Pam. richtersi group 23.01/804 Mac. richtersi 30T0437017, 4521195 15 Pam. richtersi group 23.01/807 Mac. richtersi 30T0437017, 4521195 21 Pam. richtersi group

344 ROSZKOWSKA and KACZMAREK / Turk J Zool

Table 3. (Continued).

23.01/809 Mac. richtersi 30T0437017, 4521195 11 Pam. richtersi group 23.01/814 Mac. richtersi 30T0437017, 4521195 2 Pam. richtersi group 23.01/819 Mac. richtersi 30T0437017, 4521195 1 Pam. richtersi group 23.01/820 Mac. richtersi 30T0437017, 4521195 17 Pam. richtersi group 23.01/822 Mac. richtersi 30T0437017, 4521195 16 Pam. richtersi group 23.01/823 Mac. richtersi 30T0437017, 4521195 16 Pam. richtersi group 23.01/825 Mac. richtersi 30T0437017, 4521195 8 Pam. richtersi group 23.01/829 Mac. richtersi 30T0437017, 4521195 5 Pam. richtersi group 23.01/831 Mac. richtersi 30T0437017, 4521195 17 Pam. richtersi group 23.01/832 Mac. richtersi 30T0437017, 4521195 15 Pam. richtersi group 23.01/836 Mac. richtersi 30T0437017, 4521195 5 Pam. richtersi group 23.01/837 Mac. richtersi 30T0437017, 4521195 15 Pam. richtersi group 23.01/839 Mac. richtersi 30T0437017, 4521195 16 Pam. richtersi group 23.01/843 Mac. richtersi 30T0437017, 4521195 1 Pam. richtersi group 23.01/852 Mac. richtersi 30T0435161, 4520796 6 Pam. richtersi group 23.01/869 Mac. richtersi 30T0447569, 4506108 1 Pam. sklodowskae 23.01/870 Mac. richtersi 30T0447569, 4506108 16 Pam. sklodowskae 23.01/871 Mac. richtersi 30T0447569, 4506108 17 Pam. sklodowskae 23.01/872 Mac. richtersi 30T0447569, 4506108 13 Pam. richtersi group 23.01/346 Mil. tardigradum 30T0435667, 4533269 1 Mil. beasleyi 23.01/874 Mac. richtersi 30T0447569, 4506108 11 Pam. richtersi group 23.01/875 Mac. richtersi 30T0447569, 4506108 9 Pam. richtersi group 23.01/888 Mac. richtersi 30T0442434, 4524168 2 Pam. richtersi group 23.01/890 Mac. richtersi 30T0441624, 4533508 11 Pam. richtersi group 23.01/893 Mac. richtersi 30T0449988, 4536504 7 Pam. richtersi group 23.01/431 Mil. tardigradum 30T0448682, 4542713 1 Mil. beasleyi 23.01/897 Mac. richtersi 30T0449988, 4536504 9 Pam. richtersi group 23.01/458 Mil. tardigradum 30T0451167, 4542845 1 Mil. beasleyi 23.01/90 Mac. areolatus 30T0419517, 4521376 4 Pam. areolatus group 23.01/900 Mac. richtersi 30T0449988, 4536504 11 Pam. richtersi group 23.01/482 Mil. tardigradum 30T0452915, 4521442 1 Mil. beasleyi 23.01/903 Mac. richtersi 30T0449988, 4536504 8 Pam. richtersi group 23.01/751 Mil. tardigradum 30T0429715, 4531202 1 Mil. beasleyi 23.01/909 Mac. richtersi 30T0449988, 4536504 8 Pam. richtersi group 23.01/755 Mil. tardigradum 30T0429195, 4530946 1 Mil. beasleyi 23.01/911 Mac. richtersi 30T0449988, 4536504 2 Pam. richtersi group 23.01/780 Mil. tardigradum 30T0437885, 4512612 1 Mil. beasleyi 23.01/915 Mac. richtersi 30T0449988, 4536504 3 Pam. richtersi group 23.01/873 Mil. tardigradum 30T0447569, 4506108 1 Mil. beasleyi 23.01/92 Mac. areolatus 30T0419678, 4522186 9 Pam. areolatus group 23.01/921 Mac. richtersi 30T0441091, 4537401 4 Pam. richtersi group 23.01/927 Mac. richtersi 30T0441091, 4537401 2 Pam. richtersi group 23.01/936 Mac. richtersi 30T0448433, 4548409 16 Pam. richtersi group

345 ROSZKOWSKA and KACZMAREK / Turk J Zool

Table 3. (Continued).

23.01/94 Mac. areolatus 30T0419678, 4522186 12 Pam. areolatus group 23.01/941 Mac. richtersi 30T0448433, 4548409 13 Pam. richtersi group 23.01/946 Mac. richtersi 30T0448433, 4548409 11 Pam. richtersi group 23.01/947 Mil. tardigradum 30T0448433, 4548409 1 Milnesium sp. Cuticle sculpture not visible 23.01/948 Mac. richtersi 30T0448433, 4548409 13 Pam. richtersi group 23.01/950 Mac. richtersi 30T0448433, 4548409 1 Pam. richtersi group 23.01/952 Mac. richtersi 30T0448433, 4548409 8 Pam. richtersi group 23.01/953 Mil. tardigradum 30T0448433, 4548409 1 Mil. beasleyi 23.01/956 Mac. richtersi 30T0448433, 4548409 11 Pam. richtersi group 23.01/961 Mac. richtersi 30T0448433, 4548409 9 Pam. richtersi group 23.01/97 Mac. richtersi 30T0421842, 4507133 15 Pam. richtersi group 23.01/98 Mac. richtersi 30T0421842, 4507133 1 Pam. richtersi group 23.01/740 Mil. tardigradum 30T0423927, 4521102 1 Mil. cf. dornensis Male many samples eggs were not found. When the eggs were the specimen corresponded perfectly to the original present, Mac. richtersi specimens were attributed to Pam. description. Of the above-mentioned species, the following sklodowskae Michalczyk, Kaczmarek & Węglarska, 2006 are new records to the Spanish fauna: Mil. berladnicorum, or Pam. cf. sklodowskae, if eggs were damaged and not all Mil. beasleyi, Mil. dornensis, and P. sklodowskae (for exact structures were visible or if some measurements differed localities see Table 3). On the other hand, Mac. recens, in some eggs or individuals, while morphologically they Mes. harmsworthi, Mil. eurystomum, Mil. tardigradum, were identical with original species description. Milnesium Mil. tetralamellatum, and Pam. richtersi are now doubtful tardigradum was not confirmed on any of examined slides. records and the presence of these species in Spain needs Specimens with claw configuration [3–3]–[3–3] were confirmation. attributed mainly to Mil. dornensis Ciobanu, Roszkowska Nomenclatural acts: This work and the nomenclatural & Kaczmarek, 2015 or Mil. cf. dornensis (if some acts it contains have been registered in ZooBank. measurements were different in comparison to the original The ZooBank Life Science Identifier (LSID) for this description), whereas those with claw configuration publication is: http://zoobank.org/urn:lsid:zoobank. [2–3]–[3–2] also had clearly visible pseudopores on the org:pub:CE377A13-2BC5-4B5D-8C00-2548BD212C8A cuticle and were attributed to Mil. beasleyi Kaczmarek, Jakubowska & Michalczyk, 2012. We decided to classify Acknowledgements several individuals as Mil. cf. beatae because of some This research received support from the SYNTHESYS differences in measurements in comparison to the Project (http://www.synthesys.info/), which is financed original description as well as the fact that Mil. beatae by European Community Research Infrastructure Action Roszkowska, Ostrowska & Kaczmarek, 2015 has so far under the FP7 “Capacities” Programme at the Museo been reported only from Argentina (type locality) and the Nacional de Ciencias Naturales (CSIC) or Real Jardín United States (Roszkowska et al., 2015; Tibbs et al., 2016). Botánico (CSIC) (ES-TAF-6499). Studies have been Milnesium berladnicorum Ciobanu, Zawierucha, Moglan partially conducted in the framework of activities of the & Kaczmarek, 2014 was reported only on one slide, but BARg (Biodiversity and Astrobiology Research group).

References

Bąkowski M, Roszkowska M, Gawlak M, Kaczmarek Ł (2016). Bertolani R, Guidetti R, Marchioro T, Altiero T, Rebecchi L et al. Macrobiotus naskreckii sp. nov. a new tardigrade (Eutardigrada: (2014). Phylogeny of Eutardigrada: new molecular data and Macrobiotidae: hufelandi group) from Mozambique. Annales their morphological support lead to the identification of new Zoologici 66 (2): 155-164. doi: 10.3161/00034541ANZ2016.6 evolutionary lineages. Molecular Phylogenetics and Evolution 6.2.001 76: 110-126. doi: 10.1016/j.ympev.2014.03.006 Beasley CW (1995). The phylum Tardigrada. Third Edition by G. Bertolani R, Rebecchi L (1993). A revision of the Macrobiotus Ramazzotti and W. Maucci, English Translation. Published by hufelandi group (Tardigrada, Macrobiotidae), with some the Translator. Abilene, TX, USA: Clark Beasley. observations on the taxonomic characters of . Zoologica Scripta 22: 127-152. doi: 10.1111/j.1463-6409.1993. tb00347.x

346 ROSZKOWSKA and KACZMAREK / Turk J Zool

Biserov VI (1990a). On the revision of the genus Macrobiotus. The Kaczmarek Ł, Jakubowska N, Michalczyk Ł (2012). Current subgenus Macrobiotus s. str.: a new systematic status of the knowledge on Turkish tardigrades with a description group hufelandi (Tardigrada, Macrobiotidae). Communication of Milnesium beasleyi sp. nov. (Eutardigrada: Apochela: I. Zoologicheskii Zhurnal 69: 5-17 (in Russian). Milnesiidae, the granulatum group). Zootaxa 3589: 49-64. Biserov VI (1990b). On the revision of the genus Macrobiotus. The Kaczmarek Ł, Michalczyk Ł (2017). The Macrobiotus hufelandi subgenus Macrobiotus s. str.: a new systematic status of the group (Tardigrada) revisited. Zootaxa 4363 (1): 101-123. doi: hufelandi group (Tardigrada, Macrobiotidae). Communication 10.11646/zootaxa.4363.1.4 II. Zoologicheskii Zhurnal 69: 38-50 (in Russian). Marley NJ, McInnes SJ, Sands CJ (2011). Phylum Tardigrada: a Ciobanu DA, Roszkowska M, Kaczmarek Ł (2015). Two new re-evaluation of the Parachela. Zootaxa 2819: 51-64. doi: tardigrade species from Romania (Eutardigrada: Milnesiidae, 10.11646/zootaxa2819.1.2 Macrobiotidae), with some remarks on secondary sex characters in Milnesium dornensis sp. nov. Zootaxa 3941 (4): Maucci W (1979). I Pseudechiniscus del gruppo cornutus con 542-564. doi: 10.11646/zootaxa.3941.4.4 descrizione di una nuova specie (Tardigrada, Echiniscidae). Zeszyty Naukwe Uniwersytetu Jagiellońskiego, Prace Ciobanu DA, Zawierucha K, Moglan I, Kaczmarek Ł (2014). Zoologiczne Kraków 25: 107-124 (in Italian). Milnesium berladnicorum sp. n. (Eutardigrada, Apochela, Milnesiidae), a new species of water bear from Romania. Maucci W (1991). Tre nouve specie di Eutardigradi della Groenlandia ZooKeys 429: 1-11. doi: 10.3897/zookeys.429.7755 Meridionale. Bollettino del Museo Civico di Storia Naturale di Verona 15: 279-289 (in Italian). Cuénot L (1932). Tardigrades. In: Lechevalier P (editor). Faune de France. Paris, France: Fédération Française des Sociétés de McInnes SJ (1994). Zoogeographic distribution of terrestrial/ Sciences Naturelles, pp. 1-96 (in French). freshwater tardigrades from current literature. Journal of Natural History 28: 257-352. doi: 10.1080/00222939400770131 Degma P, Guidetti R (2007). Notes to the current checklist of Tardigrada. Zootaxa 1579: 41-53. Michalczyk Ł, Kaczmarek Ł (2003). A description of the new tardigrade Macrobiotus reinhardti (Eutardigrada: Doyère LMF (1840). Mémoire sur les Tardigrades. I. Annales des Macrobiotidae, harmsworthi group) with some remarks on the Sciences Naturelles, Paris, Series 2 14: 269-362 (in French). oral cavity armature within the genus Macrobiotus Schultze. Durante Pasa MV, Maucci W (1979). Moss Tardigrada from the Zootaxa 331: 1-24. doi: 10.11646/zootaxa.331.1.1 Scandinavian Peninsula. Zeszyty Naukwe Uniwersytetu Jagiellońskiego, Prace Zoologiczne Kraków 79 (25): 47-85. Michalczyk Ł, Kaczmarek Ł (2013). The Tardigrada Register: a comprehensive online data repository for tardigrade taxonomy. Guidetti R, Bertolani R (2005). Tardigrade taxonomy: an updated Journal of Limnology 72 (S1): 175-181. doi: 10.4081/ check list of the taxa and a list of characters for their jlimnol.2013.s1.e22 identification. Zootaxa 845: 1-46. doi: 10.11646/zootaxa.845.1.1 Michalczyk Ł, Kaczmarek Ł, Węglarska B (2006). Macrobiotus Guidetti R, Schill RO, Bertolani R, Dandekar T, Wolf M (2009). sklodowskae sp. nov. (Tardigrada: Eutardigrada: Macrobiotidae, New molecular data for tardigrade phylogeny, with the richtersi group) from Cyprus. Zootaxa 1371: 45-56. doi: erection of Paramacrobiotus gen. nov. Journal of Zoological 10.11646/zootaxa.1371.1.4 Systematics and Evolutionary Research 47 (4): 315-321. doi: 10.1111/j.1439-0469.2009.00526.x Michalczyk Ł, Wełnicz W, Frohme M, Kaczmarek Ł (2012a). Redescriptions of three Milnesium Doyère, 1840 taxa Guil N (2002) Diversity and distribution of Tardigrades (Bilateria, (Tardigrada: Eutardigrada: Milnesiidae), including the Tardigrada) from the Iberian .Peninsula, Balearic Islands and nominal species for the genus. Zootaxa 3154: 1-20. doi: Chafarinas Islands. Graellsia 58: 75-94. 10.11646/zootaxa.3154.1.1 Guil N, Sanchez-Moreno S (2013). Fine-scale patterns in micrometazoans: tardigrade diversity, community composition Michalczyk Ł, Wełnicz W, Frohme M, Kaczmarek Ł (2012b). and trophic dynamics in leaf litter. Systematics and Biodiversity Corrigenda of Zootaxa, 3154: 1-20 Redescriptions of three 11 (2): 181-193. doi: 10.1080/14772000.2013.798370 Milnesium Doyère, 1840 taxa (Tardigrada: Eutardigrada: Milnesiidae), including the nominal species for the genus. Guil N, Sánchez-Moreno S, Machordom A (2009). Local biodiversity Zootaxa 3393: 66-68. doi: 10.11646/zootaxa.3393.1.6 patterns in micrometazoans: are tardigrades everywhere? Systematics and Biodiversity 7: 259-268. Morek W, Gąsiorek P, Stec D, Blagden B, Michalczyk Ł (2016). Experimental taxonomy exposes ontogenetic variability Kaczmarek Ł, Cytan J, Zawierucha K, Diduszko D, Michalczyk and elucidates the taxonomic value of claw configuration in Ł (2014). Tardigrades from Peru (South America), with Milnesium Doyère, 1840 (Tardigrada: Eutardigrada: Apochela). descriptions of three new species of Parachela. Zootaxa 3790 Contributions to Zoology 85 (2): 173-200. (2): 357-379. doi: 10.11646/zootaxa.3790.2.5 Murray J (1907). Arctic Tardigrada, collected by Wm. S. Bruce. Kaczmarek Ł, Gawlak M, Bartels PJ, Nelson DR, Roszkowska M Transactions of the Royal Society of Edinburgh 45: 669-681. (2017). Revision of the genus Paramacrobiotus Guidetti et al., doi: 10.1017/s0080456800011789. 2009 with the description of a new species, re-descriptions and a key. Annales Zoologici 67: 627-656. doi: 10.3161/00034541A Murray J (1911). Arctiscoida. Proceedings of the Royal Irish NZ2017.67.4.001 Academy 31: 1-16.

347 ROSZKOWSKA and KACZMAREK / Turk J Zool

Nelson DR, Guidetti R, Rebecchi L (2015). Phylum Tardigrada. In: Schultze CAS (1834). Macrobiotus Hufelandii animal e crustaceorum Thorp J, Rogers DC (editors). Ecology and General Biology. classe novum, reviviscendi post diuturnam asphixiam et Thorp and Covich’s Freshwater Invertebrates. London, UK: aridiatem potens. Berolini Apud Carolum Curths 1834: 1-8 (in Academic Press, pp. 347-380. doi: 10.1016/b978-0-12-385026- Latin). 3.00017-6 Schultze CAS (1840). Echiniscus bellermanni, animal crustaceum. Perry E, Miller WR, Kaczmarek Ł (2019). Recommended Scribebam Berolini 1-8 (in Latin). abbreviations for the names of genera of the phylum Tardigrada. Schuster RO, Nelson DR, Grigarick AA, Christenberry D Zootaxa (in press). (1980). Systematic criteria of Eutardigrada. Transactions Pilato G (1981). Analisi di nuovi caratteri nello studio degli of the American Microscopical Society 99: 284-303. doi: Eutardigradi. Animalia 8: 51–57 (in Italian). 10.2307/3226004 Pilato G, Bertolani R (2004). Macrobiotus dariae sp. n., a new species Stec D, Krzywański Ł, Michalczyk Ł (2018). Integrative description of eutardigrade (Eutardigrada, Macrobiotidae) from Cyprus. of Macrobiotus canaricus sp. nov. with notes on M. recens Zootaxa 638: 1-7. doi: 10.11646/zootaxa.638.1.1 (Eutardigrada: Macrobiotidae). European Journal of Taxonomy 452: 1-36. doi: 10.5852/ejt.2018.452 Pilato G, Binda MG (1991). Milnesium tetralamellatum new species of Milnesiidae from Africa (Eutardigrada). Tropical Zoology 4: Thulin G (1928). Über die Phylogenie und das System der 103-106. doi: 10.1080/03946975.1991.10539480 Tardigraden. Hereditas 11: 207-266 (in German). doi: 10.1111/ j.1601-5223.1928.tb02488.x Pilato G, Binda MG (2010). Definition of families, subfamilies, genera and subgenera of the Eutardigrada, and keys to Tibbs L, Emanuels E, Miller WR (2016). Tardigrades of the canopy: their identification. Zootaxa 2404: 1-52. doi: 10.11646/ Argentine species Milnesium beatae Roszkowska, Ostrowska zootaxa.2404.1.1 and Kaczmarek, 2015 (Eutardigrada, Milnesidae) discovered in the trees of Kansas, U.S.A. Transactions of the Kansas Academy Ramazzotti G, Maucci W (1983). Il Phylum Tardigrada. Memorie of Science 119:173-178. doi: 10.1660/062.119.0207 dell’Istituto Italiano di Idrobiologia. Verbania, Italy: Istituto Italiano di Idrobiologia (in Italian). Vecchi M, Cesari M, Bertolani R, Jönsson IK, Rebecchi L et al. (2016). Integrative systematic studies on tardigrades from Antarctica Richters F (1926). Tardigrada. In: Kükenthalund W, Krumbach T identify new genera and new species within Macrobiotoidea (editors). Handbuch der Zoologie, Vol 3. Berlin, Germany: De and Echiniscoidea. Invertebrate Systematics 30: 303-322. Gruyter, pp. 1-68 (in German). Roszkowska M, Stec D, Ciobanu DA, Kaczmarek Ł (2016). Tardigrades from Nahuel Huapi National Park (Argentina, South America) with descriptions of two new Macrobiotidae species. Zootaxa 4105: 243-260. doi: 10.11646/zootaxa.4105.3.2.

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