Bull 859 Bull. Iraq nat. Hist. Mus. (2025) 18 (4): 859-873. https://doi.org/10.26842/binhm.7.2025.18.4.0859 ORIGINAL ARTICLE PRELIMINARY DATA ON TARDIGRADA FROM THE ARMENIAN FORESTED AREA Anushik Khachatryan*, Oleg Shcherbakov** and Noushig Zarikian* *Department of Invertebrate Ecology, Scientific Center of Zoology and Hydroecology, National Academy of Sciences of Armenia, Armenia **Laboratory of Molecular Parasitology, Scientific Center of Zoology and Hydroecology, National Academy of Sciences of Armenia, Armenia Corresponding author: noushigz@hotmail.com Received: 23 June 2025, Revised: 11 Sep. 2025, Accepted: 14 Sep. 2025, Published:20 December 2025 This work is licensed under a Creative Commons Attribution 4.0 International License ABSTRACT Tardigrada is a phylum that has never been studied in Armenia, along with many other arthropod groups. Our research is about terrestrial tardigrade species. This study is the first paper on tardigrades from Armenia. Six species of tardigrade were identified, belonging to five genera: Macrobiotus hufelandi C.A.S. Schultze, 1834; Paramacrobiotus areolatus (Murray, 1907); Paramacrobiotus richtersi (Murray, 1911); Ramazzottius cf. oberhaeuseri (Doyère, 1840); Milnesium tardigradum Doyère, 1840; and Echiniscus testudo (Doyère, 1840), which are present in lichen and moss from the northern forests of Tavush Province in Armenia. All the mentioned are new to the fauna of Armenia. Keywords: Armenia, Forest diversity , New records, Tardigrada, Water bears. INTRODUCTION Tardigrades inhabit diverse ecosystems, ranging from terrestrial environments such as mosses and lichens to the deepest abysses of the oceans (Nelson et al., 2015). Notably, their ability to withstand extreme conditions, including desiccation, high levels of radiation, and even the vacuum of space, has elevated them to the status of extremophiles (Jönsson, 2007; Møbjerg et al., 2011). These remarkable adaptations challenge our understanding of the limits of life on Earth and have led researchers to explore the potential implications for astrobiology, biotechnology, and environmental science (Jönsson et al., 2008). Early taxonomic studies in the Caucasus region have identified Tardigrada species belonging to genera such as Echiniscus C. A. S. Schultze, 1840, and Milnesium Doyère, 1840 (Ramazzotti and Maucci, 1983; Dastych, 1988). However, the overall diversity and distribution patterns remain poorly explored. Recent advancements in molecular techniques and imaging technologies offer unprecedented opportunities to unveil the hidden biodiversity BULLETIN OF THE IRAQ NATURAL HISTORY MUSEUM Iraq Natural History Research Center & Museum, University of Baghdad https://jnhm.uobaghdad.edu.iq/index.php/BINHM/Home Copyright © Bulletin of the Iraq Natural History Museum Online ISSN: 2311-9799, Print ISSN: 1017-8678 https://doi.org/10.26842/binhm.7.2025.18.4.0859 https://orcid.org/0009-0003-7731-8711 https://orcid.org/0000-0001-7533-1670 https://orcid.org/0000-0001-8334-8413 mailto:noushigz@hotmail.com https://creativecommons.org/licenses/by/4.0/ https://jnhm.uobaghdad.edu.iq/index.php/BINHM/Home 860 Bull. Iraq nat. Hist. Mus. 18 (4): 859-873. Preliminary data on Tardigrada of Tardigrada in this region (Topstad et al., 2021), but the Armenian fauna of Tardigrada remains unexplored . Armenia, nestled at the crossroads of Western Asia and Eastern Europe, boasts a diverse topography ranging from high mountainous terrains to lowland areas, creating a mosaic of ecosystems. Preliminary studies have hinted at the presence of Tardigrada in Armenian environments, but a comprehensive understanding of their diversity and ecological roles within the region is yet to be established. Only one species belonging to aquatic Tardigrada has been described from Sevan Lake Pseudobiotus megalonyx (Thulin, 1928) (Krilov et al., 2010). The interdisciplinary nature of Tardigrada research is evident in its relevance to fields such as astrobiology, where the study of extremophiles informs our understanding of potential life beyond Earth (Sloan, 2018). So, this paper seeks to bridge this knowledge gap by offering a comprehensive examination of the Tardigrada fauna in Armenia, shedding light on their taxonomy, distribution, and potential ecological significance. MATERIALS AND METHODS Sampling and specimens examined: Nine sites in the mountainous forested area of Armenia were sampled in 2022-2023. Geographic coordinates, temperature during the collection season, and elevation are provided in Table (1) and Map (1). The flora of these forests includes diverse species, mainly Acer campestre L., Celtis caucasica Willd., Carpinus orientalis Mill., Elaeagnus angustifolia L., Fagus orientalis Lipsky, Fraxinus excelsior L., Quercus macranthera Fisch. & C. A. Mey. ex Hohen, Juglans regia L., Juniperus foetidissima Willd., Pistacia mutica F &M, Pinus kochiana Klotzsch ex K. Koch, Populus tremula L., Salix caprea L., Taxus baccata L., and Tilia cordata Mill . The specimens were examined in the experimental zoology laboratory of the Scientific Center of Hydroecology and Zoology of NAS RA using a Levenhook D95 LCD digital microscope and a Bresser trinocular microscope, and examined for terrestrial tardigrades, using a standard method described by Dastyc (1980). Identification: The species were identified based on keys mentioned in many references: Bertolani (1971), Dewel and Clark (1973), Dewel and Wallis (1973), Bertolani and Mambrini (1977), Ramazzotti and Maucci (1983), Biserov (1990, 1998), Bertolani and Rebecchi (1993), Biserov and Trumanov (1993), Guidetti and Bertolani (2005), Guidetti et al. (2005, 2013, 2019, 2021), Horikawa et al. (2006), Bertolani et al. (2010, 2011), Wełnicz et al. (2011), Kaczmarek et al. (2012, 2015), Vicente and Bertolani (2013), Suzuki (2016, 2022), Kaczmarek and Michalczyk (2017), Fontoura et al. (2017), Stec et al. (2018, 2020), Morek et al. (2019), and Kayastha et al. (2023a, b). The material is kept in the Scientific Center of Hydroecology and Zoology of NAS RA invertebrate collection as preparations. 861 BULLETIN OF THE IRAQ NATURAL HISTORY MUSEUM Khachatryan et al. Map (1): The locality of recorded species of Tardigrada in Armenia (by Simple mapper). Table (1): The study area of Tardigrada. Site No. Site name Coordinates Altitude m.a.s.l The highest temperature of collected day ̊ C 1 Dilijan 40.769095 N, 44.922545 E 1200 +13 2 Karaberd 40.837661 N, 44.553144 E 2400 +18 3 Kapan 41.094901 N, 44.711738 E 922 +24 4 Pambak 40.82788 N, 44.55829 E 1250 +22 5 Vanadzor 40.815524 N, 44.546165 E 1240 +23 6 Zangezur National Park 39.152721 N, 46.089392 E 2050 +19 7 Tatev 39.38611 N, 46.24451 E 1503 +35 8 Trchkan 40.915187 N, 44.114010 E 1860 +4 9 Aghveran 40.4903 N, 44.6001 E 1720 +24 RESULTS AND DISCUSSION The current study documented six species belonging to five genera. These species of tardigrade will be shown as follows: Class: Eutardigrada Order: Parachela Family: Macrobiotidae Genus Macrobiotus C.A.S. Schultze, 1834 Macrobiotus hufelandi C.A.S. Schultze, 1834 (Pl. 1) Synonym: Macrobiotus schultzei Greef, 1866 Material examined: Syunik Prov., Zangezur National Park, 26. vii.2023; Tavush Prov., Dilijan, 09.x.2022. Distribution: Worldwide (Kaczmarek and Michalczyk, 2019). 862 Bull. Iraq nat. Hist. Mus. 18 (4): 859-873. Preliminary data on Tardigrada Habitat in Armenia: This species has been recorded from soil and moss. Description of adults: Light-brown body with eye spots. Body length: 510 μm; cuticle with small pores; buccal armature characterized by an anterior and posterior band of well-defined teeth, followed dorsally and ventrally by three distinct transverse crests. Ventral median crest rounded; two pillar-shaped macroplacoids and microplacoid, the first macroplacoid longer than the second. Legs have infrequent granulation located behind the claws on fourth pair, limited and located laterally on the first three legs. Y-shaped claws, with conspicuous accessory points on the main branch; small lunulas with smooth margins present on the first three pairs of legs; larger on the fourth pair. Comments: There are no certain records for this species in the Caucasus or neighboring Iran, but it has been recorded in Turkey (Berdi et al., 2024). This is the first record for Armenia and the region. Genus Paramacrobiotus Guidetti, Schill, Bertolani, Dandekar and Wolf, 2009 Paramacrobiotus areolatus (Murray, 1907) (Pl. 2) Synonym: Macrobiotus areolatus Murray Material examined: Tavush Prov., Dilijan, 09.x.2022. Distribution: Worldwide distribution (Kayastha et al., 2023 a, b). Habitat in Armenia: This species was recorded from forest mosses and other moist habitats, where P. richtersi was found too. Description of the adults. This species has a white body. Body length 450 μm, without microplacoid; a thin cuticular thickening is present in its place, and granulations are situated on I-IV pair of legs. Eyes present, lunules under the claws of IV, and three rows of macroplacoids visible. Body of cuticle smooth, with the number of rows in the oral cavity armature ranging from I to III. Comments: This species was not recorded in Iran nor in the Caucasus, but it was recorded in Turkey (Berdi and Altındag, 2020), and is the first record for the fauna of Armenia. Paramacrobiotus richtersi (Murray, 1911) (Pl. 3) Synonym: Macrobiotus richtersi Murray, 1911 Material examined: Tavush Prov., Dilijan, 09.x.2022, Distribution: The species has a worldwide distribution (Guidetti et al., 2019). Habitat in Armenia: This species is primarily found in humid environments, especially in forest trees. Description of the adults: body brownish with length 300 μm, microplacoid exists, fine granulations situated on the I-III pair of legs, eyes absent, lunules under claws IV smooth, dorsal cuticle covered with small polygons; pt values of macroplacoid more than 48; number of rows in the oral cavity armature I-III. 863 BULLETIN OF THE IRAQ NATURAL HISTORY MUSEUM Khachatryan et al. Comments: This species was recorded in the neighboring country (Turkey) (Kaczmarek et al., 2012), but this is the first record for the fauna of Armenia. Family: Ramazzottiidae Genus Ramazzottius Binda & Pilato, 1987 Ramazzottius cf. oberhaeuseri (Doyère, 1840) (Pl. 4) Synonym: Ramazzottius oberhaeuseri (Doyère, 1840) Material examined: Lori Prov., Karaberd, 26.iii.2023 Distribution: Cosmopolitan (Stec et al., 2018). Habitat in Armenia: This species was recorded from mosses and lichens in sparse forests north of Armenia. Description of the adults: Body with red-brown colored and length 490 μm, microplacoid absent, inner claws short and thick. maroon granules are arranged in nine horizontal bands, featuring visible longitudinal rows devoid of pigment. Eyes absent, and hind claws with small and smooth pseudolunules, but more developed under claw IV. Two separated macroplacoids are visible; the body cuticle is smooth; oral cavity armature consists of two posterior bands of teeth. Comments. This species was recorded in Turkey (Kaczmarek et al., 2012) but not in the Caucasus, and is the first record for the fauna of Armenia. Order: Apochela Family: Milnesiidae Genus Milnesium Doyère, 1840 Milnesium tardigradum Doyère, 1840 (Pl. 4) Synonyms: Arcrophanes schlagintweitii Ehrenberg, 1859 Arctiscon tardigradum Schrank, 1803 Material examined: Lori Prov., Karaberd, 26.iii.2023. Distribution: Cosmopolitan (Brotto-Guidetti et al., 2024). Habitat in Armenia: This species was recorded from mosses of forested areas in Armenia. Description of the adults: Body with brownish transparent, Body length 926 μm, black eyes present, and pharyngeal bulb elongated without placoids. Placoids absent, basal spur on outer claws present with rounded basal thickenings. body cuticle without pores. Comments: This species was not recorded in neighboring countries and is the first record for the Armenian fauna. Class: Heterotardigrada Order: Echiniscoidea Family: Echiniscidae Genus Echiniscus Schultze, 1840 Echiniscus testudo (Doyère, 1840) (Pl. 6) 864 Bull. Iraq nat. Hist. Mus. 18 (4): 859-873. Preliminary data on Tardigrada Synonyms: Echiniscus bellermanni C.A.S. Schultze, 1840 Echiniscus biunguis C.A.S. Schultze, 1861 Echiniscus filamentosus Plate, 1888 Echiniscus glaber Bartoš, 1937 Echiniscus inermis Richters, 1902 Echiniscus peruvianus Binda & Pilato, 1994 Emydium testudo Doyère, 1840 Echiniscus trifilis Rahm, 1925 Material examined: Lori Prov., Karaberd, 26.iii.2023. Distribution: Europe, Africa, Asia, N. America and S. America (Kaczmarek et al., 2015, 2016). Habitat in Armenia: This species was recorded from mosses and lichens in sparse forests north of Armenia. Description of the adults: Body with orange-colored and dorsal spines present. Body length 442 μm. Microplacoid absent. Eyes present and red, and all internal claws with recurved spurs, and all external claws simple; macroplacoids absent. Body cuticle composed of polygons and light granulation on all legs. Body appendage configurations A, B, C, D, E are visible. Comments: This species was recorded in Turkey and Iran (Berdi and Altında, 2020; Mokhtari et al., 2018) and wasn’t recorded in the Caucasus countries. It is the first record for the fauna of Armenia. Forest cover, comprising 11.2% of Armenia, represents one of the country's less-studied ecosystems. However, with a growing focus on climate change mitigation, research in these forests is expanding, highlighting all components of their biodiversity. Given the mountainous terrain of Armenia's forested regions, access is challenging, and field conditions are suitable only during a brief period in summer. Consequently, findings on Tardigrada in these areas remain preliminary. All species included in this study represent the first records for Armenian fauna. Six species Macrobiotus hufelandi, Paramacrobiotus richtersi, Paramacrobiotus areolatus, Ramazzottius oberhaeuseri, Echiniscus testudo, and Milnesium tardigradum were previously unknown from both the Caucasus and the Armenian Highland, likely due to the scarcity of studies in this region and the historical neglect of Tardigrada research here. Armenia’s forest biodiversity, particularly that of Tardigrades, is believed to be quite rich, yet the results from preliminary research are insufficient to clarify their distribution and diversity. Future studies may reveal more about the distribution, traits, and potentially even new species of these fascinating organisms. 865 BULLETIN OF THE IRAQ NATURAL HISTORY MUSEUM Khachatryan et al. Plate (1): Macrobiotus hufelandi; (A) General view, (B) Buccal apparatus, (C) Claws, (D) Egg. Plate (2): Paramacrobiotus areolatus; (A) General view, (B) Buccal apparatus, (C) Claws, (D) Egg. 866 Bull. Iraq nat. Hist. Mus. 18 (4): 859-873. Preliminary data on Tardigrada Plate (3): Paramacrobiotus richtersi; (A) General view, (B) Buccal apparatus, (C) Claws, (D) Egg. Plate (4): Ramazzottius cf. oberhaeuseri; (A) General view, (B) Claws, (C) Buccal apparatus, (D) Buccal apparatus. 867 BULLETIN OF THE IRAQ NATURAL HISTORY MUSEUM Khachatryan et al. Plate (5): Milnesium tardigradum; (A) General view, (B) Buccal apparatus, (C) Eggs, (D) Claws. Plate (6): Echiniscus testudo; (A) General view, (B) Cuticle, (C) Claws, (D) Buccal apparatus, (E) Eggs. CONCLUSIONS This study presents the first comprehensive record of forest-dwelling Tardigrada species in Armenia, documenting six species Macrobiotus hufelandi, Paramacrobiotus richtersi, 868 Bull. Iraq nat. Hist. Mus. 18 (4): 859-873. Preliminary data on Tardigrada Paramacrobiotus areolatus, Ramazzottius cf. oberhaeuseri, Milnesium tardigradum, and Echiniscus testudo none of which had previously been reported from Armenia, the Caucasus, or the Armenian Highland. These findings significantly expand the known distribution ranges of these taxa and highlight the underexplored nature of Armenia’s forest ecosystems. Given the challenging field conditions and limited seasonal window for research, the data presented here should be regarded as preliminary. Nevertheless, the discovery of these cosmopolitan and widespread taxa in Armenia underscores the potential richness of tardigrade diversity in the region. The mountainous forest habitats of Armenia, though covering a modest portion of the country, appear to harbor a distinct and potentially diverse tardigrade fauna . Although this study does not include quantitative measures of abundance, diversity indices, or statistical analyses correlating species occurrence with environmental variables, this was a deliberate methodological choice given its baseline, exploratory nature. Our primary aim was to document, for the first time, the terrestrial tardigrade fauna of Armenia through accurate morphological identification and distributional reporting. For regions and taxa with no prior terrestrial records, such faunistic inventories are an essential first step before more complex quantitative analyses can be meaningfully applied. The environmental metadata we provide— including precise coordinates, altitudes, and microhabitat details—form a foundational dataset that will enable future studies to incorporate multi-season sampling, standardized abundance assessments, diversity metrics, and ecological modeling. ACKNOWLEDGMENTS This research was supported by funding from the Science Committee of the Ministry of Education, Science, Culture, and Sports of the Republic of Armenia with grant code 24AA- F011 . CONFLICT OF INTEREST STATEMENT "The authors declare that there is no conflict of interest". LITERATURE CITED Berdi, D. and Altindag, A. 2020. Contribution to the knowledge on distribution of Tardigrada in Turkey. Diversity, 12(3): 95. [CrossRef] Berdi, D., Nassouhi, D., Çiftci, E., Güner, B., Trampacı, B., Gülsoy, S. and Altındağ, A. 2024. Acute toxicity of lead on the survival of Macrobiotus hufelandi (Eutardigrada: Parachela: Macrobiotidae). Journal of Wildlife and Biodiversity, 8(2): 38-54. 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(2025) 18 (4): 859-873. من املناطق الغابية في أرمينيا Tardigradaبينات أولية عن بطيئات الخطو زاريكيان* كنوش و أنوشيك خاتشاتريان*، أوليغ شيرباكوف** * قسم علم بيئة الالفقاريات، املركز العلمي لعلم الحيوان والبيئة املائية، األكاديمية ، أرمينياالوطنية للعلوم في أرمينيا ** مختبر علم الطفيليات الجزيئية، املركز العلمي لعلم الحيوان والبيئة املائية، األكاديمية ، أرمينياالوطنية للعلوم في أرمينيا 20/12/2025، النشر: 14/9/2025القبول: ،11/9/2025املراجعة: ،23/6/2025االستالم: الخالصة الخطو البطيئات عد ُ إلى جانب Tardigradaت أرمينيا، في درس قط ُ ت لم التي عب ُ الش من البرية. البطيئات الخطو أنواع ز بحثنا على ّ يرك األخرى. املفصليات العديد من مجموعات عد هذه الدراسة أول عمل علمي يتناول البطيئات الخطو في أرمينيا. تم تحديد ستة أنواع ُ وت من هذه الكائنات، تنتمي إلى خمسة أجناس وهي: Macrobiotus hufelandi (C.A.S. Schultze, 1834( Paramacrobiotus areolatus (Murray, 1907 ( Paramacrobiotus richtersi (Murray, 1911 ( Ramazzottius cf. oberhaeuseri (Doyère, 1840 ( Milnesium tardigradum (Doyère, 1840( Echiniscus testudo (Doyère, 1840 ( تم العثور عليها في األشنيات والطحالب املوجودة في غابات مقاطعة تافوش الشمالية في عد جميع هذه األنواع جديدة على عالم االحياء األرمنية. ُ أرمينيا. وت