Vol.:(0123456789) Organisms Diversity & Evolution (2025) 25:167–187 https://doi.org/10.1007/s13127-024-00668-5 ORIGINAL ARTICLE Checklist of tardigrades in Germany as a contribution to biodiversity research Rolf Schuster1 · Ralph O. Schill2 Received: 24 September 2024 / Accepted: 2 December 2024 / Published online: 4 February 2025 © The Author(s) 2025 Abstract Over 250 years ago Johann August Ephraim Goeze, a Protestant pastor from Quedlinburg, Germany, discovered the first tardigrade. The German physiologist and anatomist Carl August Sigismund Schultze (1795–1877) later named the first spe- cies Macrobiotus hufelandi C.A.S. Schultze, 1834, a designation that remains valid today. By the first third of the twentieth century numerous new species had been discovered in Germany and were comprehensively compiled for the first time by the German zoologist Ernst Marcus in 1936. Since then additional faunistic studies have been conducted. This new checklist of tardigrades in Germany provides an overview of all known species found in the country to date. It includes 91 limno- terrestrial or limnic species and eight marine species, with 21 belonging to Heterotardigrada and 78 to Eutardigrada. Germany is the type locality (terra typica) for 24 tardigrade species. The number of identified tardigrade species varies significantly depending on the extent of studies conducted in different federal states. Baden-Württemberg has the highest number of spe- cies identified, with 80 species recorded across eleven studies. In this state the Black Forest, with its remarkable diversity of 72 identified tardigrade species is one of the most intensively studied regions worldwide. In Hesse 30 species have been recorded from five studies, while Berlin has documented 23 species from two studies. Fewer species have been reported from other federal states. The 99 tardigrade species identified in Germany represent about 7% of the total 1,488 described tardigrade species worldwide. Keywords Tardigrada · Germany · Checklist · Taxonomy · Literature · Biodiversity · Microinvertebrates Introduction The animal group of tardigrades was discovered in Ger- many over 250 years ago. Johann August Ephraim Goeze (1731–1793) was a Protestant pastor in Quedlinburg, Ger- many (Saxony-Anhalt, ST). He began his career as a trans- lator of various scientific and philosophical works and only in the last third of his life became a highly respected and productive zoologist, particularly as a helminthologist. He was the first to describe a tardigrade, probably a Hypsibius species. This description was published under the title “Von einigen merkwürdigen Wasserinsekten” (On some strange water insects) in an appendix of the German translation of Bonnet’s "Traité d’Insectologie." The appendix contains per- sonal observations made by J.A.E. Goeze in the environment of his hometown Quedlinburg. He discovered water bears in duckweed (especially in winter) and wrote: “Seltsam ist dieses Thierchen, weil der ganze Bau seines Körpers ausse- rordentlich und seltsam ist, und weil es in seiner äusserlichen Gestalt, dem ersten Anblicke nach, die größte Aehnlichkeit mit einem Bäre im Kleinen hat. Das hat mich auch bewogen, ihm den Namen des kleinen Wasserbärs zu geben”. (Strange is this little creature, because the whole organisation of its body is extraordinary and strange and because of its external appearance. At the first glance, it has the closest similarity to a little bear. This also led me to give it the name little water bear.) (Goeze, 1773). In 1834 Carl August Sigismund Schultze (1795–1877), a German physiologist and anatomist, named the first species Macrobiotus hufelandi C.A.S. Schultze, 1834a, 1834b, which is still valid today (Degma & Guidetti, 2024). Between 1927 * Ralph O. Schill ralph.schill@bio.uni-stuttgart.de Rolf Schuster r.b.schuster@web.de 1 Rolf Schuster, Mattenweg 9, 79856 Hinterzarten, Germany 2 University of Stuttgart, Institute of Biomaterials and biomolecular Systems, Pfaffenwaldring 57, 70569 Stuttgart, Germany http://orcid.org/0000-0002-7930-5282 http://crossmark.crossref.org/dialog/?doi=10.1007/s13127-024-00668-5&domain=pdf 168 R. Schuster, R. O. Schill and 1936 the German zoologist Ernst Marcus published four substantial and wide-ranging articles (Marcus, 1927, 1928, 1929, 1936). These included a synopsis of all tardigrade spe- cies previously described around the world in the series “Die Tierwelt Deutschlands” (Fauna of Germany). This first and as yet only overview of tardigrades in Germany (within the borders of 1918) was published nearly one hundred years ago (Marcus, 1928, 1936). However no other checklist of tardi- grade species found in Germany has been compiled since. Marcus (1936) reported 44 species (valid according to Degma & Guidetti, 2024) from Germany. By this study, we aim to provide an overview of the tardigrade species found in Ger- many from the discovery and description of the first species to the present day. 169Checklist of tardigrades in Germany as a contribution to biodiversity research Materials and methods The basis for this current checklist is the first summary of German tardigrades by Marcus (1928, 1936) and the data from Ramazzotti and Maucci (1983). The sources cited by Marcus primarily refer to the authors, and the listed tardigrade locations were assigned to the 16 federal states of Germany. Only those locations from Marcus that lie within the present-day territory of the Federal Republic of Germany were included. The term "Germany" refers to the borders of present-day Germany. The unspecific locality "Northern Germany" from Marcus (1936) encompasses the federal states Brandenburg (BB), Berlin (BE), Bremen (HB), Hamburg (HB), Mecklenburg-Vorpommern (MV), Lower Saxony (NI) and Schleswig–Holstein (SL); "Central Germany" North Rhine-Westphalia (NW), Saxony-Anhalt (ST), Saxony (SN) and Thuringia (TH); and "Southern Germany" refers to Baden-Württemberg (BW), Bavaria (BY), Rhineland-Palatinate (RP) and Saarland (SL). The term "German Central Uplands" includes the federal states of Central and Southern Germany. Rahm (1925) studied tardigrades in the Rhine Province, a former administrative unit, which is now part of NW and RP. The term "Harz" in Marcus (1936) refers to the federal states NI, ST and TH. All faunistic and taxonomic studies on tardigrades conducted in Germany after Marcus's compilation (1936) have been incorporated into the current checklist of tar- digrades in Germany. This includes recently discovered species from the last few years (Guidetti et  al., 2021, 2022a; Morek et al., 2019; Schuster, 2021). Species used for physiological, molecular and biophysical studies which also originate from Germany are included in the current checklist of tardigrades of Germany but are not marked on the map of study areas (Fig. 1) (e.g., Baumann, 1961; Greven & Peters, 1986). Animal cultures of some species were established and subsequently used for various stud- ies and publications over longer periods. Since these ani- mals can be attributed to an original location and therefore do not represent a new site, these publications were only considered once. The species gathered from the literature were checked against the worldwide checklist with valid species names (Degma & Guidetti, 2024). Invalid and syn- onymized species were also listed (Tables 1, 2). Species for which Germany is the terra typica were highlighted separately (Table 3). Also included in the list (Table 4) and on the map (Fig. 1) is the species Milnesium tardigradum Doyère, 1840. To account for the significantly expanded Milnesium tax- onomy in recent years, a redescription was carried out by Michalczyk et al. (2012). The neotype of M. tardigradum s.s. and its 46 neoparatypes now originate from Zeesen, (BB), Germany. Currently, there is no comprehensive study of tardigrades across all of Germany. Most research has been concentrated in specific regions: Richters examined the tardigrade fauna around Frankfurt (Richters, 1902) in Hesse and along the Mecklenburg coast (Richters, 1904a) in Mecklenburg-Vor- pommern. Heinis conducted studies in the southern Black Forest (Heinis, 1910) in Baden-Württemberg, while Rahm (1925) focused on the Rhine Province, an area that now includes parts of North Rhine-Westphalia and Rhineland- Palatinate. The current checklist of tardigrades of Germany, summarizing all species found up until 1936, was compiled by Marcus (1936). After Marcus emigrated to Brazil in 1936 (Greven, 2018) following his dismissal as Associate Professor from the Frie- drich Wilhelm University of Berlin due to his Jewish faith, only a few studies on the distribution of tardigrades in Ger- many were conducted. Englisch (1936) investigated areas around Berlin, Hesse-Nassau and a region in the Harz Moun- tains, while Schultz (1935, 1951, 1953) focused on marine tardigrades from the North and Baltic Seas. Baumann (1961, 1964, 1966, 1970) reported on individual tardigrade species in Bremen and Lower Saxony. Fig. 1 Study areas on tardigrades in Germany. The areas where more than five species were identified are marked in red. Areas where a new species was described are marked with an asterisk (*). (1) Schultze (1834a, 1834b) near St. Ulrich, near Freiburg (BW); (2) Plate (1888) near Marburg (HE); (3) Richters (1900, 1902, 1903) in Frankfurt and surroundings (Taunus), (HE); (4) Richters (1904a) at Heiligendamm, Sassnitz, Markgrafenheide, Müritz (MV); (5) Rich- ters (1907a) at Ludwigshafen (RP); (6) Richters (1908) at the Kiel Fjord (SH); (7) Richters (1909a) in the Eckernförde Bay (SH); (8) Heinis (1910) in the Southern Black Forest (BW); (9) Rahm (1925) at Rhineland Province (now parts of NW and RP), further surround- ings of Bonn (NW); (10) Marcus (1928) at Schlachtensee (BE); (11) Schulz (1935) at Helgoland (SH); (12) Marcus (1936) in the Harz (SH); (13) Marcus (1936) near Wartburg, near Eisenach (TH); (14) Englisch (1936) near Berlin and surroundings, Harz (Brocken) (ST), Hesse-Nassau, near Marburg (HE); (15) Schulz (1951) at Sylt (SH); (16) Schulz (1953) at Amrum (SH); (17) Greven (1972) in the northern Sauerland near Volkringshausen (NW); (18) Adler (1987) at the University of Stuttgart-Hohenheim (BW); (19) Hofmann and Eichelberg (1986) in the Lahnauen near Gießen (HE); (20) Berto- lani and Rebecchi (1993) near St. Ulrich, near Freiburg (BW); (21) Russel et al. (1994) in the Sandhausen Dunes near Heidelberg (BW); (22) Grabowski (1995) in the Lahnbergen near Marburg (HE); (23) Schuster (1999) near Hinterzarten, near Freiburg (BW); (24) Schuster (2003a, 2003b) in Hinterzarten and surroundings (BW); (25) Hohberg (2006) at Berzdorf, near Dresden (SN); (26) Hohberg et  al. (2011) at Welzow, near Spremberg (BB); (27) Michalczyk et  al. (2012) in Zeesen (BB); (28) Morek et al. (2019) in Bebenhausen, near Tübin- gen (BW); (29) Bingemer et  al. (2020) at Welzow, near Spremberg (BB); (30) Guidetti et  al. (2021) in the Black Forest National Park (BW); (31) Schuster (2021) in the Höllental (Black Forest), near Freiburg (BW); (32) Guidetti et  al. (2022a) in the Black Forest National Park (BW). Abbreviations: BB = Brandenburg, BE = Berlin, BW = Baden-Württemberg, BY = Bavaria, HB = Bremen, HE = Hesse, HH = Hamburg, MV = Mecklenburg-Vorpommern, NI = Lower Saxony, NW = North Rhine-Westphalia, RP = Rhineland-Palatinate, SH = Schleswig–Holstein, SL = Saarland, SN = Saxony, ST = Saxony- Anhalt, TH = Thuringia ◂ 170 R. Schuster, R. O. Schill Ta bl e 1 S pe ci es w ith u nc er ta in ta xo no m ic v al ue re po rte d fro m G er m an y Sp ec ie s N am e Fe de ra l S ta te Re fe re nc e St at us M ac ro bi ot us m ac ro ny x D uj ar di ni , 1 85 1, D ac ty lo bi ot us m ac ro ny x (D uj ar di n, 1 85 1) BW , H E, N W R ic ht er s, 19 09 b; H ei ni s, 19 10 ; M ar cu s, 19 36 sp ec ie s d ub ia e t i nq ui re nd a ac cc or di ng to D as ty ch (1 98 8) , n om en d ub iu m a cc or di ng to D as ty ch (2 01 5) an d Po gw iz d an d St ec (2 02 0) D ip ha sc on a lp in um (M ur ra y, 1 90 6a ), H yp si bi us (D ip ha sc on ) a lp in us (M ur ra y, 1 90 6a ) B E, H E M ar cu s, 19 36 ; E ng lis ch , 1 93 6; H of m an n & E ic he l- be rg , 1 98 6; A dl er , 1 98 7; H of m an n, 1 98 7 no m en d ub iu m a cc or di ng to D as ty ch (2 01 5) a nd Tu m an ov a nd T sv et ko va (2 02 3) D ip ha sc on c hi le ne ns e Pl at e, 1 88 8, H yp si bi us (D ip ha s- co n) c hi le ne ns is P la te , 1 88 8 H E R ic ht er s, 19 04 b; M ar cu s, 19 36 no m en d ub iu m a cc or di ng to D as ty ch (2 01 5) Ec hi ni sc us a rc to m ys E hr en be rg , 1 85 3 BW , N W , R P R ic ht er s, 19 02 , 1 90 4a ; H ei ni s, 19 10 ; R ah m , 1 92 5; M ar cu s, 19 36 no m en in qu ire nd um a cc or di ng to G ąs io re k et  a l. (2 01 9a ) Ec hi ni sc us m us ci co la P la te , 1 88 8 BW , H E, N W , R P R ic ht er s, 19 02 ; H ei ni s, 19 10 ; R ah m , 1 92 5 sp ec ie s i nq ui re nd a ac co rd in g to M ar cu s ( 19 36 ) Ec hi ni sc us si m ili s P la te , 1 88 8 H E, N W , S N , S T, T H Pl at e, 1 88 8; M ar cu s, 19 28 sp ec ie s d ub ia e t i nq ui re nd a ac co rd in g to M ar cu s ( 19 36 ) H yp si bi us w ib be lti R ah m , 1 92 5 N W , R P R ah m , 1 92 5 sp ec ie s d ub ia e t i nq ui re nd a ac co rd in g to M ar cu s ( 19 36 ) Is oh yp si bi us p ap ill ife r b ul bo su s ( M ar cu s, 19 28 ), H yp si bi us p ap ill ife r f . b ul bo su s ( M ar cu s, 19 28 ), H yp si bi us (I so hy ps ib iu s) p ap ill ife r f or m a bu lb os a (M ar cu s, 19 28 ) B E M ar cu s, 19 36 su pp re ss ed b y G ąs io re k et  a l. (2 01 9b ) Is oh yp si bi us sc ha ud in ni (R ic ht er s, 19 09 b) , M ac ro - bi ot us sc ha ud in ni (R ic ht er s, 19 09 b) , H yp si bi us (I so hy ps ib iu s) sc ha ud in ni (R ic ht er s, 19 09 b) B E, H E, N W , S T R ic ht er s, 19 09 b; M ar cu s, 19 36 ; E ng lis ch , 1 93 6; H of - m an n & E ic he lb er g, 1 98 6; H of m an n, 1 98 7 no m en in qu ire nd um a cc or di ng to G ąs io re k et  a l. (2 01 9b ) Is oh yp si bi us tu be rc ul at us (P la te , 1 88 8) , M ac ro bi ot us tu be rc ul at us (P la te , 1 88 8) , H yp si bi us (I so hy ps ib iu s) tu be rc ul at us (P la te , 1 88 8) B E, B W , H E Pl at e, 1 88 8; M ar cu s, 19 28 , 1 93 6; H of m an n & E ic he l- be rg , 1 98 6; A dl er , 1 98 7 no m en d ub iu m a cc or di ng to D as ty ch (2 01 5) ; n om en in qu ire nd um a cc or di ng to G ąs io re k et  a l. (2 01 9b ) M ac ro bi ot us fe rd in an di R eu ka uf , 1 91 2a H E, N W , S N , S T, T H Re uk au f, 19 12 a; M ar cu s, 19 36 R am az zo tti a nd M au cc i ( 19 83 ) d es cr ib ed th e or ig in al id en tifi ca tio n pr oc es s a s n ea rly im po ss ib le O re el la v ilu ce ns is R ah rm , 1 93 1 N W R ah m , 1 93 1; M ar cu s, 19 36 ; R am az zo tti & M au cc i, 19 83 sp ec ie s i nq ui re nd a ac co rd in g to D eg m a an d G ui de tti (2 02 4) M ac ro bi ot us h ib er ni cu s ( M ur ra y, 1 91 1) , H yp si bi us hi be rn ic us (M ur ra y, 1 91 1) BW Sc hu ste r, 20 03 a, 2 00 3b no m en d ub iu m a cc or di ng to G ui de tti e t a l. (2 02 2b ) Ra m az zo tti us n ov em ci nc tu s ( M ar cu s, 19 36 ), H yp si bi us (H yp si bi us ) n ov em ci nc tu s ( M ar cu s, 19 36 ) N W , R P M ar cu s, 19 36 no m en d ub iu m a cc or di ng to S te c et  a l. (2 01 8) U rs ul in iu s n od os us (M ur ra y, 1 90 7d ), M ac ro bi ot us no do su s ( M ur ra y, 1 90 7d ), H yp si bi us (I so hy ps ib iu s) no do su s ( M ur ra y, 1 90 7d ) B E M ar cu s, 19 28 , 1 93 6 no m en d ub iu m a cc or di ng to D as ty ch (2 01 5) 171Checklist of tardigrades in Germany as a contribution to biodiversity research Ta bl e 2 S pe ci es re po rte d fro m G er m an y un de r s yn on ym ou s s pe ci es n am es Va lid S pe ci es N am e Fe de ra l S ta te Pr ev io us S pe ci es N am e Re fe re nc e A ut ho rit y C al oh yp si bi us o rn at us (R ic ht er s, 19 00 ) BW , B Y, H E, N W , R P, S L, S N , S T, TH H yp si bi us (C al oh yp si bi us ) o rn at us fo rm a sp in io si ss im a R ic ht er s, 19 00 M ar cu s, 19 36 sy no ny m iz ed w ith C al oh yp si bi us o rn at us ac co rd in g to R am az zo tti & M au cc i, 19 83 Ec hi ni sc us b lu m i R ic ht er s, 19 03 H E Ec hi ni sc us b is et os us H ei ni s, 19 08 H of m an n & E ic he lb er g, 1 98 6 sy no ny m iz ed w ith E ch in is cu s b lu m i ac co rd in g to G ąs io re k et  a l., 2 01 9b Ec hi ni sc us m er ok en si s R ic ht er s, 19 04 a, 19 04 b B B Ec hi ni sc us m er ok en si s s ue ci cu s T hu lin , 19 11 M ar cu s, 19 36 sy no ny m iz ed w ith E ch in is cu s m er ok en si s ac co rd in g to G ąs io re k & V on či na , 2 02 3 Ec hi ni sc us q ua dr is pi no su s R ic ht er s, 19 02 H E Ec hi ni sc us sc ro fa R ic ht er s, 19 02 R ic ht er s, 19 02 ; M ar cu s, 19 36 sy no ny m iz ed w ith E ch in is cu s q ua dr is - pi no su s a cc or di ng to R am az zo tti & M au cc i, 19 83 Ec hi ni sc us te st ud o (D oy èr e, 1 84 0) H E, N W , R P Ec hi ni sc us fi la m en to su s P la te , 1 88 8 R ic ht er s, 19 02 ; R ah m , 1 92 5; M ar cu s, 19 36 sy no ny m iz ed w ith E ch in is cu s t es tu do ac co rd in g to G ąs io re k et  a l., 2 01 7 Ec hi ni sc us te st ud o (D oy èr e, 1 84 0) H E Ec hi ni sc us in er m is R ic ht er s, 19 02 R ic ht er s, 19 02 ; M ar cu s, 19 36 sy no ny m iz ed w ith E ch in is cu s t es tu do ac co rd in g to M ar cu s, 19 36 Ec hi ni sc us te st ud o (D oy èr e, 1 84 0) N W , R P Ec hi ni sc us tr ifi lis R ah m , 1 92 4 R ah m , 1 92 5; M ar cu s, 19 36 sy no ny m iz ed w ith E ch in is cu s t es tu do ac co rd in g to M ar cu s, 19 36 H yp si bi us c on ve rg en s ( U rb an ow ic z, 19 25 ) BW M ac ro bi ot us te tra da ct yl us G re ef , 1 86 6 H ei ni s, 19 10 ; M ar cu s, 19 36 sy no ny m iz ed w ith H yp si bi us c on ve rg en s ac co rd in g to M ar cu s, 19 36 G re ve ni us a sp er (M ur ra y, 1 90 6c ) N I, N W , R P, T H , S T H yp si bi us te tra da ct yl oi de s ( R ic ht er s, 19 07 b) ; H yp si bi us (I so hy ps ib iu s) te tra - da ct yl oi de s ( R ic ht er s, 19 07 b) R ah m , 1 92 5; M ar cu s, 19 36 sy no ny m iz ed w ith G re ve ni us a sp er ac co rd in g to D as ty ch , 2 01 6 Th ul in iu s a ug us ti (M ur ra y, 1 90 7c ) BW M ac ro bi ot us la cu st ri s D uj ar di n, 1 85 1; M ac ro bi ot us a ug us ti M ur ra y, 1 90 7a , 19 07 b, 1 90 7c , 1 90 7d ; H ys ib iu s (I so hy si bi us ) a ug us ti M ur ra y, 1 90 7a , 19 07 b, 1 90 7c , 1 90 7d R ic ht er s, 19 09 b; H ei ni s, 19 10 ; W en ck , 19 14 sy no ny m iz ed w ith T hu lin iu s a ug us ti by G ąs io re k et  a l., 2 01 9b H yp si bi us d uj ar di ni (D oy èr e, 1 84 0) Th M ac ro bi ot us la cu st ri s D uj ar di n, 1 85 1 Re uk au f, 19 12 c sy no ny m iz ed w ith H yp si bi us d uj ar di ni ac co rd in g to M ar cu s, 19 36 O rz el is cu s b el op us d u B oi s- Re ym on d M ar cu s, 19 52 SH O rz el is cu s s ep te nt ri on al is S ch ul z, 1 95 3 Sc hu lz , 1 95 3 sy no ny m iz ed w ith O rz el is cu s b el op us ac co rd in g to P ol lo ck , 1 98 2 172 R. Schuster, R. O. Schill From the 1970s onward smaller areas began to be studied for tardigrades. Greven (1972) explored tardigrades in the northern Sauerland region (NW), Adler (1987) conducted research at the University of Hohenheim in Stuttgart (BW) and Hofmann and Eichelberg (1986) studied the fauna near Gießen (HE). Russel et al. (1994) focused on tardigrades in sand dunes near Heidelberg (BW). In 1995 Grabowski examined tardigrades in mosses from the Lahnbergen Hills near Marburg (HE), and in 2003, Schuster investigated tardi- grades in the southern Black Forest (BW). Soil-dwelling tar- digrades were later studied by Hohberg (2006) near Berzdorf (SN), by Hohberg et al. (2011) and by Bingemer et al. (2020) near Welzow (BB). Michalczyk et al. (2012) studied tardi- grades in Zeesen (BB). Since 2016, the tardigrade fauna of the Black Forest National Park in the Northern Black Forest (BW) has been studied as part of a large-scale biodiversity assessment and monitoring program (pers. communication R. Schill & R. Guidetti). Results & Discussion In 1936 Marcus reported 60 tardigrade species for the area of present-day Germany. The tardigrade species listed by Marcus from the former eastern territories of Germany (now Poland) are not included in this list. Of these 60 spe- cies Marcus classified three as species dubia et inquirenda (Table 1). Of the remaining 57 species, 44 are still listed in the current checklist (Degma & Guidetti, 2024). 15 species identified in Germany are now considered either nomen dubium (e.g. Diphascon alpinum), species dubia et inquirenda (e.g. Echiniscus similis), or have insuffi- cient original descriptions (e.g. Macrobiotus ferdinandi) (Table 1). Twelve species have been synonymized with previously identified species (e.g. Echiniscus scrofa is syn- onymous with Echiniscus quadrispinosus; Table 2). According to the current checklist with valid scien- tific names (Degma & Guidetti, 2024) a total of 99 tardi- grade species have been recorded in Germany, including 91 limno-terrestrial or limnic species and eight marine species (Table 4). Of these 99 species, 21 belong to Het- erotardigrada and 78 to the class Eutardigrada (Table 4). The 99 species represent about 7% of the total described tardigrade species of 1,488 worldwide (Degma & Guidetti, 2024). In similarly sized European countries, the number of tardigrade species is comparable: Norway (excluding Svalbard) has 102 species (Meier, 2017), Sweden and Poland each have 101 species (Guidetti et al., 2015), while Finland has fewer species 68 (Vuori et al., 2020) and Ire- land has 51 (DeMilio et al., 2016). However, Italy has a Table 3 Tardigrade species described from Germany and their terra typica. The numbers in the table refer to the overview map (Fig. 1) Species Terra typica No Actinarctus doryphorus doryphorus Schulz, 1935 Helgoland (SH) 11 Batillipes mirus Richters, 1909a Eckernförder Bay (SH) 7 Bryocherus intermedius laevis (Marcus, 1936) Wartburg near Eisenach (TH) 13 Calohypsibius ornatus (Richters, 1900) Taunus (HE) 3 Cornechiniscus cornutus (Richters, 1907a) near Ludwigshafen (RP) 5 Crenubiotus ruhesteini Guidetti et al., 2021 Black Forest (BW) 29 Diphascon birklehofi Rolf Schuster, 1999 Black Forest (BW) 23 Dianea sattleri (Richters, 1902) Taunus (HE) 3 Diphascon pingue pingue (Marcus, 1936) Wartburg (TH) 12 Doryphoribius evelinae (Marcus, 1928) Schlachtensee Berlin (BE) 10 Echiniscus blumi Richters, 1903 Taunus (HE) 3 Echiniscus quadrispinosus Richters, 1902 Taunus (HE) 3 Fractonotus verrucosus (Richters, 1900) Taunus (HE) 3 Halobiotus stenostomus (Richters, 1908) Kieler Fjord (SH) 6 Macrobiotus echinogenitus Richters, 1903 Taunus (HE) 3 Macrobiotus hufelandi C.A.S. Schultze, 1834a, 1834b St. Ulrich near Freiburg (BW) 1 Macrobiotus sandrae Bertolani & Rebecchi, 1993 St. Ulrich near Freiburg (BW) 20 Milnesium tardigradum Doyère, 1840 Zeesen (BB) 27 Milnesium inceptum Morek et al., 2019 Bebenhausen near Tübingen (BW) 28 Minibiotus bernhardi R. Schuster, 2021 Black Forest (BW) 30 Minibiotus intermedius (Plate, 1888) Marburg (HE) 2 Pilatobius granifer (Greven, 1972) Sauerland (NW) 17 Ramazzottius kretschmanni Guidetti et al., 2022a Black Forest (BW) 31 Stygarctus bradypus Schulz, 1951 Sylt (SH) 15 173Checklist of tardigrades in Germany as a contribution to biodiversity research Ta bl e 4 O ve rv ie w o f T ar di gr ad a sp ec ie s r ec or de d in G er m an y. N — no nm ar in e sp ec ie s ( lim no -te rr es tri al o r l im ni c on es ), M — m ar in e sp ec ie s G en us Sp ec ie s A ut ho rit y Fe de ra l S ta te M ar in e Re fe re nc e H ET ER O TA R D IG R A D A M ar cu s, 19 27 A RT H RO TA R D G R A D A M ar cu s, 19 27 B at ill ip ed id ae R am az zo tti , 19 62 1 Ba til lip es m ir us R ic ht er s, 19 09 a SH M R ic ht er s, 19 09 a; R em an e, 1 92 4; M ar cu s, 19 27 ; M ar cu s, 19 36 ; Sc hm id t, 19 71 a 2 Ba til lip es ph re at ic us Re na ud D eb ys er , 1 95 9 N I, SH M Po llo ck , 1 97 1 3 Ba til lip es tu be rn at is Po llo ck , 1 97 1 N I, SH M Po llo ck , 1 97 1; R am az zo tti & M au cc i, 19 83 H al ec hi ni sc id ae T hu lin , 1 92 8 O rz el isc in ae S ch ul z, 1 96 3 4 O rz el is cu s be lo pu s du B oi s- Re ym on d M ar cu s, 19 52 (O . s ep te nt ri on al is Sc hu lz , 1 95 3) SH M Sc hu lz , 1 95 3; R am az zo tti & M au cc i, 19 83 St yg ar ct id ae S ch ul z, 1 95 1, St yg ar ct in ae S ch ul z, 1 95 1 5 St yg ar ct us br ad yp us Sc hu lz , 1 95 1 SH M Sc hu lz , 1 95 1; S ch m id t, 19 71 a Ta na rc tid ae R en au d- M or - na nt , 1 98 0 6 Ac tin ar ct us do ry ph or us d or yp ho ru s Sc hu lz , 1 93 5 SH M Sc hu ltz , 1 93 5; G re ll, 1 93 6; M ar cu s, 19 36 EC H IN IS C O ID EA R ic ht er s, 19 26 Ec hi ni sc oi di da e K ris te ns en & H al la s, 19 80 7 Ec hi ni sc oi de s si gi sm un di (M .S ch ul tz e, 1 86 5) SH M Sc hu ltz e, 1 86 5; R ic ht er s, 19 08 ; M ar cu s, 19 27 , 1 93 6; G re ve n & G ro hé , 1 97 5; G re ve n & K ris te ns en , 2 00 1 Ec hi ni sc id ae T hu lin , 1 92 8 8 Br yo ch er us in te rm ed iu s l ae vi s (M ar cu s, 19 36 ) TH N M ar cu s, 19 36 ; R am az zo tti & M au cc i, 19 95 9 Br yo de lp ha x pa rv ul us Th ul in , 1 92 8 BY N M ar cu s, 19 36 10 Br yo de lp ha x ta tre ns is (W ęg la rs ka , 1 95 9) BY N W ęg la rs ka , 1 95 9; R am az zo tti & M au cc i, 19 95 11 C la xt on ia we nd ti (R ic ht er s, 19 03 ) BW N H ei ni s, 19 10 ; R ah m , 1 92 5; M ar cu s, 19 36 174 R. Schuster, R. O. Schill Ta bl e 4 (c on tin ue d) G en us Sp ec ie s A ut ho rit y Fe de ra l S ta te M ar in e Re fe re nc e 12 C or ne ch in is cu s co rn ut us (R ic ht er s, 19 07 a) BW , R P N R ic ht er s, 19 07 a; H ei ni s, 19 10 ; M ar cu s, 19 36 13 Ec hi ni sc us bl um i R ic ht er s, 19 03 B B , B E, B W , B Y, H E, M V, N I, N W , R P N R ic ht er s, 19 03 ; H ei ni s, 19 10 ; R ah m , 1 92 5; M ar cu s, 19 28 , 19 36 ; E ng lis ch , 1 93 6; B au - m an n, 1 97 0; A dl er , 1 98 7; H of m an n & E ic he lb er g, 1 98 6; H of m an n, 1 98 7; S ch us te r, 20 03 a, 2 00 3b 14 Ec hi ni sc us gr an ul at us (D oy èr e, 1 84 0) B B , B E, B W , H E, N W N R ic ht er s, 19 03 ; M ar cu s, 19 28 ; En gl is ch , 1 93 6; G re ve n, 1 97 2; Sc hu ste r, 20 03 a, 2 00 3b ; S ch ill & S te in br üc k, 2 00 7 15 Ec hi ni sc us m er ok en si s R ic ht er s, 19 04 b B B , B W N M ar cu s, 19 36 ; S ch us te r, 20 03 a, 20 03 b 16 Ec hi ni sc us qu ad ri sp in os us R ic ht er s, 19 02 BW , H E, M V N R ic ht er s, 19 02 , 1 90 4a ; H ei ni s, 19 10 ; M ar cu s, 19 28 ; S ch us te r, 20 03 a, 2 00 3b ; K ac zm ar ek et  a l., 2 02 2a 17 Ec hi ni sc us sp in ul os us (D oy èr e, 1 84 0) BW N Sc hu ste r ( un pu bl is he d) 18 Ec hi ni sc us te st ud o (D oy èr e, 1 84 0) B B , B W , B Y, H B , H E, M V, N I, N W , R P N H ei ni s, 19 10 ; M ar cu s, 19 28 , 19 36 ; E ng lis ch , 1 93 6; B au m an n, 1 96 4; S ch m id t, 19 71 a; G re ve n, 1 97 2; G re ve n & P et er s, 19 86 ; A dl er , 1 98 7; H of m an n & E ic he lb er g, 1 98 6; H of m an n, 1 98 7; G ra bo w sk i, 19 95 ; S ch us te r, 20 03 a, 2 00 3b ; Sc hi ll & S te in br üc k, 2 00 7 19 Ec hi ni sc us tr is et os us C ué no t, 19 32 H E N M ar cu s, 19 36 20 Ps eu de ch in is cu s su ill us (E hr en be rg , 1 85 3) B B , B E, B W , B Y, H B , H E, H H , M V, N I, N W , R P, S L, SN , S T, S H , T H N H ei ni s, 19 10 ; M ar cu s, 19 36 ; G re ve n, 1 97 2; S ch us te r, 20 03 a, 2 00 3b 21 Te ste ch in is cu s sp its be rg en si s s pi ts be rg en si s (S co ur fie ld , 1 89 7) BW , B Y, H E, N I, N W , R P, SL , S N , S T, T H N M ar cu s, 19 36 ; R am az zo tti & M au cc i, 19 83 EU TA R D IG R A D A R ic ht er s, 19 26 A PO C H EL A S ch us te r e t a l., 19 80 M iln es iid ae R am az zo tti , 1 96 2 175Checklist of tardigrades in Germany as a contribution to biodiversity research Ta bl e 4 (c on tin ue d) G en us Sp ec ie s A ut ho rit y Fe de ra l S ta te M ar in e Re fe re nc e 22 M iln es iu m in ce pt um M or ek , S uz uk i, Sc hi ll, G eo r- gi ev , Y an ko va , M ar le y & M ic ha lc zy k, 2 01 9 BW N M or ek e t a l., 2 01 9 23 M iln es iu m ta rd ig ra du m ta rd ig ra du m D oy èr e, 1 84 0 B B , B E, B W , H B , H E, M V, N I, N W , R P, S T, S H , T H N R ic ht er s, 19 04 a; H ei ni s, 19 10 ; R ah m , 1 92 5; M ar cu s, 19 28 , 1 93 6; E ng lis ch , 1 93 6; B au m an n, 1 96 4, 1 96 6, 1 97 0; Sc hm id t, 19 71 a, 1 97 1b ; G re - ve n & B lo m , 1 97 7; G re ve n & P et er s, 19 86 ; H of m an n & Ei ch el be rg , 1 98 6; H of m an n, 19 87 ; R us se l e t a l., 1 99 4; G ra bo w sk i, 19 95 ; S ch us te r, 20 03 a, 2 00 3b ; G re ve n, 2 00 7; Sc hi ll & S te in br üc k, 2 00 7; Sc hu ste r & G re ve n, 2 00 7; M ic ha lc zy k et  a l., 2 01 2 PA R AC H EL A S ch us te r e t a l., 19 80 Eo hy ps ib io id ea B er to la ni & K ris te ns en , 1 98 7 Eo hy ps ib iid ae B er to la ni & K ris te ns en , 1 98 7 24 Be rt ol an iu s we gl ar sk ae (D as ty ch , 1 97 2) BW N Sc hu ste r, 20 03 a, 2 00 3b H yp sib io id ea P ila to , 1 96 9b C al oy hy ps ib iid ae P ila to , 19 69 b 25 C al oh yp si bi us or na tu s (R ic ht er s, 19 00 ) BW , B Y, H E, N W , R P, S L, SN , S T, T H N R ic ht er s, 19 00 ; H ei ni s, 19 10 ; M ar cu s, 19 36 ; E ng lis ch , 19 36 ; S ch us te r, 20 03 a, 2 00 3b H ys ib iid ae P ila to , 1 96 9b D ip ha sc on in ae D as ty ch , 1 99 2 26 D ip ha sc on bi rk le ho fi Ro lf Sc hu ste r, 19 99 BW N Sc hu ste r, 19 99 , 2 00 3a , 2 00 3b 27 D ip ha sc on hi gg in si B in da , 1 97 1 B B , B W , S N N Sc hu ste r, 20 03 a, 2 00 3b ; H oh be rg , 2 00 6; S ch us te r & G re ve n, 2 00 7; H oh be rg e t a l., 20 11 ; B in ge m er & H oh be rg , 20 17 ; B in ge m er e t a l., 2 02 0 28 D ip ha sc on hu m ic us B er to la ni , G ui de tti & R eb ec - ch i, 19 94 BW N Sc hu ste r, 20 03 a, 2 00 3b 176 R. Schuster, R. O. Schill Ta bl e 4 (c on tin ue d) G en us Sp ec ie s A ut ho rit y Fe de ra l S ta te M ar in e Re fe re nc e 29 D ip ha sc on no bi le i (B in da , 1 96 9) B B , B W , H E, S N N Ru ss el e t a l., 1 99 4; G ra bo w sk i, 19 95 ; S ch us te r, 20 03 a, 2 00 3b ; Sc hu ste r & G re ve n, 2 00 7; B in ge m er & H oh be rg , 2 01 7; B in ge m er e t a l., 2 02 0 30 D ip ha sc on pi ng ue p in gu e (M ar cu s, 19 36 ) BW , H E, N I, SN , S T, T h N M ar cu s, 19 36 ; R us se l e t a l., 19 94 ; G ra bo w sk i, 19 95 ; Sc hu ste r, 20 03 a, 2 00 3b ; Sc hu ste r & G re ve n, 2 00 7, 20 13 ; B in ge m er & H oh be rg , 20 17 H yp sib iin ae P ila to , 1 96 9b 31 H yp si bi us co nv er ge ns (U rb an ow ic z, 1 92 5) B B , B E, B W , B Y, H E, H B , M V, N I, N W , R P, S H , S T, TH N M ar cu s, 19 28 , 1 93 6; E ng lis ch , 19 36 ; B au m an n, 1 96 1; A dl er , 19 87 ; H of m an n & E ic he lb er g, 19 86 ; H of m an n, 1 98 7; R us - se l e t a l., 1 99 4; G ra bo w sk i, 19 95 ; R am az zo tti & M au cc i, 19 83 ; S ch us te r, 20 03 a, 2 00 3b ; Sc hu ste r & G re ve n, 2 00 7; B in ge m er & H oh be rg , 2 01 7 32 H yp si bi us du ja rd in i (D oy èr e, 1 84 0) B E, B W , H E, N I, N W , S N , TH N H ei ni s, 19 10 ; M ar cu s, 19 36 ; En gl is ch , 1 93 6; A m m er - m an n, 1 96 2, 1 96 7; S ch m id t, 19 71 a, 1 97 1c ; G re ve n, 1 97 2; H of m an n & E ic he lb er g, 1 98 6; H of m an n, 1 98 7; G ra bo w sk i, 19 95 ; S ch us te r, 20 03 a, 2 00 3b ; H oh be rg , 2 00 6; S ch us te r & G re ve n, 2 00 7; B in ge m er & H oh be rg , 2 01 7 33 H yp si bi us m ic ro ps Th ul in , 1 92 8 BW N Sc hu ste r, 20 03 a, 2 00 3b 34 H yp si bi us pa lli du s Th ul in , 1 91 1 BW , H E, S N N A dl er , 1 98 7; R us se l e t a l., 1 99 4; G ra bo w sk i, 19 95 ; S ch us te r, 20 03 a, 2 00 3b ; H oh be rg , 2 00 6; Sc hu ste r & G re ve n, 2 00 7; B in ge m er & H oh be rg , 2 01 7 35 H yp si bi us pr ad el lii B er to la ni & R eb ec ch i, 19 96 BW N Sc hu ste r, 20 03 a, 2 00 3b 36 H yp si bi us sc ab ro py gu s C ué no t, 19 29 BW , B Y, H e, N W , T H , S T, SN N M ar cu s, 19 36 ; S ch us te r, 20 03 a, 20 03 b; S ch us te r & G re ve n, 20 07 ; G ui de tti e t a l., 2 02 2a 177Checklist of tardigrades in Germany as a contribution to biodiversity research Ta bl e 4 (c on tin ue d) G en us Sp ec ie s A ut ho rit y Fe de ra l S ta te M ar in e Re fe re nc e It aq ua sc on id ae R ud es cu , 19 64 37 Ad ro pi on be lg ic ae (R ic ht er s, 19 11 ) B E, B W N M ar cu s, 19 28 , 1 93 6; S ch us te r, 20 03 a, 2 00 3b 38 Ad ro pi on sc ot ic um (M ur ra y, 1 90 5b ) B E, B W , H E, N W N H ei ni s, 19 10 ; M ar cu s, 19 36 ; En gl is ch , 1 93 6; R ie m an n, 19 66 ; H of m an n & E ic he l- be rg , 1 98 6; H of m an n, 1 98 7; G ra bo w sk i, 19 95 ; S ch us te r, 20 03 a, 2 00 3b ; S ch us te r & G re ve n, 2 00 7 39 As ta tu m en tr in ac ri ae (A rc id ia co no , 1 96 2) BW , H e, N W , S N N G re ve n, 1 97 2; G ra bo w sk i, 19 95 ; S ch us te r, 20 03 a, 2 00 3b ; H oh be rg , 2 00 6; B in ge m er & H oh be rg , 2 01 7 40 G ui de tti on m od es tu m (B in da e t a l., 1 98 4) BW N Sc hu ste r, 20 03 a, 2 00 3b ; S ch us - te r & G re ve n, 2 00 7 41 G ui de tti on pr or si ro st re (T hu lin , 1 92 8) BW , H B , H E, H H , M V, N I, SH N Th ul in , 1 92 8; M ar cu s, 19 36 ; G ra bo w sk i, 19 95 ; R am az zo tti & M au cc i, 19 83 ; S ch us te r, 20 03 a, 2 00 3b 42 Ita qu as co n pl ac op ho ru m M au cc i, 19 73 BW N pe rs . c om . S ch ill & G ui de tti , 20 20 43 M es oc ri st a sp itz be rg en si s (R ic ht er s, 19 03 ) B B , B W , H E, N I, ST , T H N R ic ht er s, 19 03 ; M ar cu s, 19 36 ; En gl is ch , 1 93 6; G ra bo w sk i, 19 95 ; S ch us te r, 20 03 a, 2 00 3b ; Sc hu ste r & G re ve n, 2 00 7 44 Pl at ic ri st a an gu st at a (M ur ra y, 1 90 5a ) BW N Sc hu ste r, 20 03 a, 2 00 3b 45 Ra ri bi us pa wl ow sk ii W ęg la rs ka , 1 97 3 BW N Sc hu ste r, 20 03 a, 2 00 3b M ic ro hy ps ib iid ae P ila to , 19 98 46 M ic ro hy ps ib iu s be rt ol an ii K ris te ns en , 1 98 2 BW N Sc hu ste r, 20 03 a, 2 00 3b 47 M ic ro hy ps ib iu s tr un ca tu s Th ul in , 1 92 8 BW N Sc hu ste r, 20 03 a, 2 00 3b Pi la to bi id ae B er to la ni e t a l., 20 14 48 D eg m io n oc ul at um (M ur ra y, 1 90 6b ) ST , B W N M ar cu s, 19 36 ; E ng lis ch , 1 93 6; Sc hu ste r, 20 03 a, 2 00 3b ; Sc hu ste r & G re ve n, 2 00 7 49 Fo nt ou ri on re ca m ie ri (R ic ht er s, 19 11 ) BW N Sc hu ste r, 20 03 a, 2 00 3b 178 R. Schuster, R. O. Schill Ta bl e 4 (c on tin ue d) G en us Sp ec ie s A ut ho rit y Fe de ra l S ta te M ar in e Re fe re nc e 50 Pi la to bi us bu lla tu s (M ur ra y, 1 90 5b ) B E, B W , S N N M ar cu s, 19 36 ; S ch us te r, 20 03 a, 20 03 b; S ch us te r & G re ve n, 20 07 ; B in ge m er & H oh be rg , 20 17 51 Pi la to bi us gr an ife r (G re ve n, 1 97 2) N W N G re ve n, 1 97 2 52 Pi la to bi us ru go su s (B ar to š, 19 35 ) BW , S N N Sc hu ste r, 20 03 a, 2 00 3b ; S ch us - te r & G re ve n, 2 00 7; B in ge m er & H oh be rg , 2 01 7 R am az zo tti id ae S an ds e t a l., 20 08 53 H eb es un cu s co nj un ge ns (T hu lin , 1 91 1) BW N Sc hu ste r, 20 03 a, 2 00 3b 54 Ra m az zo tti us kr et sc hm an ni G ui de tti , C es ar i, G io va nn in i, Eb el , F ör sc hl er , R eb ec ch i & Sc hi ll, 2 02 2 BW N G ui de tti e t a l., 2 02 2a , 2 02 2b 55 Ra m az zo tti us ob er ha eu se ri (D oy èr e, 1 84 0) B B , B E, B W , B Y, H E, N I, N W , M V, R P, S H , S T N R ic ht er s, 19 04 a; H ei ni s, 19 10 ; R ah m , 1 92 5; M ar cu s, 19 28 , 1 93 6; E ng lis ch , 1 93 6; B au m an n, 1 96 6, 1 97 0; A dl er , 19 87 ; H of m an n & E ic he l- be rg , 1 98 6; H of m an n, 1 98 7; G ra bo w sk i, 19 95 ; S ch us te r, 20 03 a, 2 00 3b ; B in ge m er & H oh be rg , 2 01 7 Is oh yp sib io id ea S an ds e t a l., 20 08 D or yp ho ri bi id ae G ąs io re k et  a l., 2 01 9a , 2 01 9b 56 Ap od ib iu s co nf us us D as ty ch , 1 98 3 B B , B W , S N N H oh be rg e t a l,. 2 01 1; D ab er t et  a l., 2 01 4; B in ge m er & H oh - be rg , 2 01 7; B in ge m er e t a l., 20 20 ; S ch us te r ( un pu bl is he d) 57 D or yp ho ri bi us ev el in ae (M ar cu s, 19 28 ) B E N M ar cu s, 19 28 , 1 93 6 58 G re ve ni us as pe r (M ur ra y, 1 90 6c ) N I, N W , R P, S T, T H N M ar cu s, 19 36 59 G re ve ni us gr an ul ife r (T hu lin , 1 92 8) BW , N W , S T N En gl is ch , 1 93 6; G re ve n & B lo m , 1 97 7; S ch us te r, 20 03 a, 20 03 b 60 Ps eu do bi ot us m eg al on yx (T hu lin , 1 92 8) B B , B E, B W , N W , R P, T H N M ar cu s, 19 36 61 Th ul in iu s au gu st i (M ur ra y, 1 90 7c ) B B , B E, B W , N W , R P, T H N R ic ht er s, 19 09 b; W en ck , 1 91 4; M ar cu s, 19 36 ; G re ve n, 1 97 2, 19 76 62 Th ul in iu s ru ffo i (B er to la ni , 1 98 1) BW N Sc hu ste r, 20 03 a, 2 00 3b 179Checklist of tardigrades in Germany as a contribution to biodiversity research Ta bl e 4 (c on tin ue d) G en us Sp ec ie s A ut ho rit y Fe de ra l S ta te M ar in e Re fe re nc e H al ob io tid ae G ąs io re k et  a l., 20 19 a, 2 01 9b 63 H al ob io tu s ste no sto m us (R ic ht er s, 19 08 ) SH M R ic ht er s, 19 08 ; M ar cu s, 19 36 ; R am az zo tti & M au cc i, 19 83 H ex ap od ib iid ae C es ar i e t a l., 20 16 64 H ex ap od ib iu s m ic ro ny x Pi la to , 1 96 9a B B N B in ge m er e t a l., 2 02 0 Is oh yp sib iid ae S an ds e t a l., 20 08 65 D ia ne a pa pi lli fe ra (M ur ra y, 1 90 5a ) B E N M ar cu s, 19 36 ; R am az zo tti & M au cc i, 19 83 66 D ia ne a sa ttl er i (R ic ht er s, 19 02 ) B E, B W , H E, N W , S T, S N N R ic ht er s, 19 02 ; M ar cu s, 19 36 ; G re ve n, 1 97 2; H of m an n & Ei ch el be rg , 1 98 6; H of m an n, 19 87 ; R us se l e t a l., 1 99 4; G ra bo w sk i, 19 95 ; S ch us te r, 20 03 a, 2 00 3b 67 Er em ob io tu s al ic at ai (B in da , 1 96 9) B B , B W , B Y, S N N Ru ss el e t a l., 1 99 4; H oh be rg , 20 06 ; H oh be rg e t a l., 2 01 1; B in ge m er & H oh be rg , 2 01 7; B in ge m er e t a l., 2 02 0; S ch us - te r ( un pu bl is he d) 68 Fr ac to no tu s ve rr uc os us (R ic ht er s, 19 00 ) B E, B W , H E N R ic ht er s, 19 00 ; H ei ni s, 19 10 ; M ar cu s, 19 36 ; R am az zo tti & M au cc i, 19 83 ; S ch us te r, 20 03 a, 2 00 3b 69 Is oh yp si bi us da st yc hi Pi la to e t a l., 1 98 2 B B , H E N G ra bo w sk i, 19 95 ; B in ge m er & H oh be rg , 2 01 7; B in ge m er et  a l., 2 02 0 70 Is oh yp si bi us pr os os to m us Th ul in , 1 92 8 B E, B B , B W , H E N M ar cu s, 19 36 ; E ng lis ch , 1 93 6; A dl er , 1 98 7; R us se l e t a l., 19 94 ; G ra bo w sk i, 19 95 ; Sc hu ste r, 20 03 a, 2 00 3b ; Sc hu ste r & G re ve n, 2 00 7; G ui de tti e t a l., 2 02 2a 71 U rs ul in iu s el eg an s (B in da & P ila to , 1 97 1) BW N Sc hu ste r ( un pu bl is he d) 72 U rs ul in us lu nu la tu s (I ha ro s, 19 66 ) H E N G ra bo w sk i, 19 95 73 U rs ul in us pa pp i (I ha ro s, 19 66 ) BW N Sc hu ste r, 20 03 a, 2 00 3b ; S ch us - te r & G re ve n, 2 00 7 74 U rs ul in us ro ns is va lle i (B in da & P ila to , 1 96 9a , 19 69 b) BW N Ru ss el e t a l., 1 99 4 180 R. Schuster, R. O. Schill Ta bl e 4 (c on tin ue d) G en us Sp ec ie s A ut ho rit y Fe de ra l S ta te M ar in e Re fe re nc e M ac ro bi ot oi de a Th ul in , 1 92 8 A do ry bi ot id ae S te c, V ec ch i & M ic ha lc zy k, 2 02 0 75 C re nu bi ot us ru he ste in i G ui de tti , S ch ill , G io va nn i, M as sa , G ol do ni , E be l, Fö r- sc hl er , R eb ec ch i & C es ar i, 20 21 BW N G ui de tti e t a l., 2 02 1 M ac ro bi ot id ae T hu lin , 1 92 8 76 M ac ro bi ot us ec hi no ge ni tu s R ic ht er s, 19 03 B E, B W , H E, N W , R P N R ic ht er s, 19 03 ; H ei ni s, 19 10 ; R ah m , 1 92 5; M ar cu s, 19 28 , 19 36 77 M ac ro bi ot us hu fe la nd i C .A .S . S ch ul tz e, 1 83 4a , 18 34 b B B , B E, B W , B Y, H B , H E, H H , M V, N I, N W , R P, S L, SN , S T, S H , T H N R ic ht er s, 19 04 a; H ei ni s, 19 10 ; R ah m , 1 92 5; M ar cu s, 19 28 , 1 93 6; E ng lis ch , 1 93 6; B au m an n, 1 97 0; G re ve n, 19 72 ; G re ve n & K uh lm an n, 19 72 ; W al z, 1 97 9; G re ve n & P et er s, 19 86 ; A dl er , 1 98 7; H of m an n & E ic he lb er g, 1 98 6; H of m an n, 1 98 7; B er to la ni & R eb ec ch i, 19 93 ; R us se l et  a l., 1 99 4; G ra bo w sk i, 19 95 ; Sc hu ste r, 20 03 a, 2 00 3b ; Sc hu ste r & G re ve n, 2 00 7, 20 13 78 M ac ro bi ot us m ac ro ca lix B er to la ni & R eb ec ch i, 19 93 BW N Sc hu ste r ( un pu bl is he d) 79 M ac ro bi ot us pe rs im ili s B in da & P ila to , 1 97 2 BW N A dl er , 1 98 7 80 M ac ro bi ot us sa nd ra e B er to la ni & R eb ec ch i, 19 93 BW N B er to la ni & R eb ec ch i, 19 93 81 M ac ro bi ot us sc ot ic us St ec , M or ek , G ąs io re k, B la gd en & M ic ha lc zy k, 20 17 BW N Sc hu ste r ( un pu bl is he d) 82 M ac ro bi ot us vl ad im ir i B er to la ni e t a l., 2 01 1 BW N B er to la ni e t a l., 2 01 1 83 M es ob io tu s fu rc ig er (M ur ra y, 1 90 7a ) B E N M ar cu s, 19 28 , 1 93 6 84 M es ob io tu s ha rm sw or th i (M ur ra y, 1 90 7b ) B B , B E, B W , H E, N I, N W , R P, S N , S T, T H N M ar cu s, 19 28 , 1 93 6; E ng lis ch , 19 36 ; A dl er , 1 98 7; H of - m an n & E ic he lb er g, 1 98 6; H of m an n, 1 98 7; R us se l e t a l., 19 94 ; G ra bo w sk i, 19 95 ; Sc hu ste r, 20 03 a, 2 00 3b 85 M es ob io tu s pa tie ns (P ila to e t a l., 2 00 0) BW N Sc hu ste r & G re ve n, 2 00 7 86 M in ib io tu s be rn ha rd i R .S ch us te r, 20 21 BW N Sc hu ste r, 20 21 181Checklist of tardigrades in Germany as a contribution to biodiversity research Ta bl e 4 (c on tin ue d) G en us Sp ec ie s A ut ho rit y Fe de ra l S ta te M ar in e Re fe re nc e 87 M in ib io tu s fu rc at us (E hr en be rg , 1 85 9) BW N Sc hu ste r, 20 03 a, 2 00 3b 88 M in ib io tu s in te rm ed iu s (P la te , 1 88 8) BW , H E, M V, N W , R P, T H , SN , S T N Pl at e, 1 88 8; R ic ht er s, 19 04 a; H ei ni s, 19 10 ; M ar cu s, 19 36 ; En gl is ch , 1 93 6; G ra bo w sk i, 19 95 ; S ch us te r, 20 03 a, 2 00 3b ; Sc hu ste r & G re ve n, 2 00 7; K ac zm ar ek e t a l., 2 02 2b 89 Pa ra m ac ro bi ot us ar eo la tu s (M ur ra y, 1 90 7c ) BW , N W , R P N R ah m , 1 92 5; M ar cu s, 19 36 ; Sc hi ll & S te in br üc k, 2 00 7 90 Pa ra m ac ro bi ot us ri ch te rs i (M ur ra y, 1 91 1) B B , B E, B W , B Y, H E, S N N M ar cu s, 19 28 , 1 93 6; A dl er , 19 87 ; H of m an n & E ic he lb er g, 19 86 ; H of m an n, 1 98 7; R us - se l e t a l., 1 99 4; G ra bo w sk i, 19 95 ; S ch us te r, 20 03 a, 2 00 3b ; H oh be rg & T ra un sp ur ge r, 20 05 ; H oh be rg , 2 00 6; S ch ill & S te in br üc k, 2 00 7; S ch us te r & G re ve n, 2 00 7, 2 01 3; B in ge - m er & H oh be rg , 2 01 7 91 Si su bi ot us sp ec ta bi lis (T hu lin , 1 92 8) BW N Sc hu ste r, 20 03 a, 2 00 3b 92 Te nu ib io tu s w ill ar di (P ila to , 1 97 7) BW N Sc hu ste r ( un pu bl is he d) 93 Xe ro bi ot us xe ro ph ilu s (D as ty ch , 1 97 8) BW N Ru ss el e t a l., 1 99 4 M ur ra yi da e G ui de tti e t a l., 20 00 94 D ac ty lo bi ot us di sp ar (M ur ra y, 1 90 7a ) B B , B E, B W , S N , T H N Re uk au f, 19 12 b; M ar cu s, 19 28 , 19 36 ; S ch us te r, 20 03 a, 2 00 3b 95 M ur ra yo n ha st at us (M ur ra y, 1 90 7c ) BW N H ei ni s, 19 10 ; M ar cu s, 19 28 , 19 36 96 M ur ra yo n pu lla ri (M ur ra y, 1 90 7c ) B E, B W , N I, ST , T H N M ar cu s, 19 36 ; E ng lis ch , 1 93 6; Sc hu ste r, 20 03 a, 2 00 3b 97 Pa ra m ur ra yo n m ei er i G ui de tti , G io va nn in i, D el Pa pa , E kr em , N el so n, Re be cc hi & C es ar i, 20 22 b BW N Sc hu ste r ( un pu bl is he d) 98 Pa ra m ur ra yo n ste lla tu s (G ui de tti , 1 99 8) BW N Sc hu ste r ( un pu bl is he d) R ic ht er siu sid ae G ui de tti et  a l., 2 02 1 99 Ri ch te rs iu s co ro ni fe r (R ic ht er s, 19 03 ) N W , R P N R ah m , 1 92 5; M ar cu s, 19 36 182 R. Schuster, R. O. Schill significantly higher number of species, with 246 species reported (Guidetti et al., 2015). The map (Fig. 1) provides an overview of the regions where tardigrade studies have been conducted from the time of the description of the first tardigrade species Macrobiotus hufelandi to the present. In these regions, either new species have been discovered and described, or more than five differ- ent tardigrade species have been found. Some regions men- tioned in the literature could be localized precisely, while vague geographical references, such as "Northern Germany" or "Central Uplands," do not feature on the overview map (Fig. 1). However, as previously mentioned, these areas have been assigned to specific federal states, ensuring that these species are also included in the current checklist. The number of identified tardigrade species varies greatly depending on the extent of studies conducted in different federal states. Baden-Württemberg (BW) has the highest number of species identified (78 species from eleven stud- ies), followed by Hesse (HE) (30 species from five studies) and Berlin (BE) (23 species from two studies). The high number of studies in Schleswig–Holstein (SH) (ten studies) with relatively few identified species is due to the marine tardigrade research, where often only one species was identi- fied (Schulz, 1935, 1951, 1953). The relatively high number of species identified in BE with only two studies is due to Marcus's intensive research in his local area. The high num- ber of species in BW is attributed to Schuster's work (2003), which identified 54 species in his local area in the southern Black Forest, in conjunction with Heinis's work (1910). It is likely that the number of tardigrade species in many federal states is higher, especially in areas with similar geographical and climatic conditions to BW. All species from these 28 studies are listed in Fig. 1 and Table 4. As in other countries, marine tardigrades have been inadequately studied. To date only eight marine tardigrade species have been identified along the coast of the North and Baltic Seas. Given the diverse ecological conditions in these seas, it is likely that additional species remain to be discovered. Similarly marine tardigrade species are under- represented in the checklists of neighboring countries of the North and Baltic Seas, with the exception of the Faroe Islands, where Hansen et al. (2001) conducted a detailed study of interstitial meiofauna and identified 35 marine tar- digrade species. Macrobiotus hufelandi is the first described tardigrade species (Schultze, 1834a, 1834b) from near Freiburg (BW) and its name is still valid today. Marcus (1936) reported its distribution across northern, central and southern Germany, covering the entire country. A revision of the Macrobio- tus hufelandi group by Bertolani and Rebecchi (1993) in the type locality of Macrobiotus hufelandi (St. Ulrich, near Freiburg im Breisgau, BW) revealed the presence of at least one additional species (Macrobiotus sandrae) very similar to Macrobiotus hufelandi. Bertolani et al. (2011) found a third species in this area, Macrobiotus vladimiri. Most of the studies mentioned above identified Macrobiotus hufelandi. It is likely that not all studies actually identified Macrobiotus hufelandi sensu stricto, particularly since many similar spe- cies were described later by Bertolani and Rebecchi (1993) and subsequently. It is expected that with an integrative taxo- nomic approach (i.e. combining morphological traits and molecular analysis for species identification), additional spe- cies in the Macrobiotus hufelandi group, previously identi- fied as such, may be discovered in future research. The taxonomy of tardigrades has changed significantly over the past two decades. Most of the listed studies date back 20 years or more, and tardigrade identification up until Bertolani et al. (2011) was based solely on morphological characteristics. With the advent of integrated descriptions of tardigrades, species identification has become increasingly differentiated and precise. Many species identifications from the past two centuries are incomplete by the standards of modern species descriptions. As a result, numerous iden- tifications must be regarded as doubtful in the context of contemporary taxonomy. This is particularly true for species such as Macrobiotus hufelandi, Pseudechiniscus suillus, Mil- nesium tardigradum, Ramazzottius oberhäuseri, Mesobiotus harmsworthi, Hypsibius convergens, Testechiniscus spits- bergensis spitsbergensis, Diphascon scoticum, and others (see Kaczmarek et al., 2014). These identifications should be reassessed in light of modern taxonomic standards. The type locality for 24 tardigrade species occurs in Ger- many (Fig. 1, Table 3). Most of these species were described in the first four decades of the past century, primarily by Richters (1900, 1902, 1903, 1907a, 1908, 1909a), Marcus (1928, 1936) and Schulz (1935). New species from Ger- many have been increasingly described in recent years: Mil- nesium inceptum in 2019, Crenubiotus ruhesteini in 2021, Minibiotus bernhardi and Ramazzottius kretschmanni in 2022 (Table 3). Kaczmarek et al., (2022a, b) redescribed Minibiotus intermedius and Echiniscus quadrispinosus using an integrative approach, and Michalczyk et al. (2012) made a redescription of Milnesium Doyère, 1840 taxa (Tardigrada: Eutardigrada: Milnesiidae), including the nominal species for the genus. Schuster (2003a, b) reported 54 species in the Black For- est, including five species identified by Heinis (1910) that were not found by Schuster. Additionally Schuster identi- fied Hypsibius cf. convergens, Isohypsibius cf. annulatus, and Ramazzottius cf. cataphractus. Since then, the follow- ing changes have occurred: Minibiotus cf. poricinctus has been described as Minibiotus bernhardi and Minibiotus cf. scopulus has been identified as Macrobiotus scoticus. Macrobiotus hibernicus is now considered nomen dubium (Degma & Guidetti, 2024). The species previously identified as Macrobiotus hibernicus are now split into Paramurrayon 183Checklist of tardigrades in Germany as a contribution to biodiversity research meieri and Paramurrayon stellatus. The species previously identified as Macrobiotus crenulatus is now Crenubiotus ruhesteini. Thanks to the work of Heinis (1910), Bertolani and Rebecchi (1993), Bertolani et al. (2011), and Guidetti et al., (2021, 2022a), all conducted in the Black Forest, this region is now the most extensively studied in Germany with 70 identified tardigrade species and it ranks among the most intensively studied regions for tardigrades worldwide. Acknowledgements We would like to express our deepest gratitude to Hartmut Greven from the University of Düsseldorf and to Franz Brüm- mer from the University of Stuttgart, both of whom actively supported and guided the authors' work over a long period. Many colleagues assisted us in research and in the compilation of this current checklist of tardigrades in Germany, and we extend our thanks to them as well. Moreover, we want to thank the anonymous reviewer, whose feedback greatly improved the quality of this manuscript. Funding Open Access funding enabled and organized by Projekt DEAL. Declarations Competing Interests The authors declare no competing interests. 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