534 Northeastern Naturalist Vol. 18, No. 4 534 The Terrestrial and Freshwater Tardigrada of Northeastern North America, with New Records from Maine Harry A. Meyer* Abstract - Tardigrada (water bear) is a phylum of microscopic animals commonly found in mosses, lichens, leaf litter, and freshwater. Over 200 species of water bear are known to occur in North America; 99 species have been reported from northeastern North America. In lichen and leaf-litter samples from Bar Harbor, Hancock County, ME (a state for which only one species of tardigrade, Thulinius augusti, has hitherto been reported), eleven species of water bear were present: Echiniscus merokensis, Milnesium tardigradum, Ramazzottius baumanni, R. oberhaeuseri, Diphascon (Diphascon) nodulosum, D. (Adropion) prorsirostre, Astatumen trinacriae, Macrobiotus harmsworthi, Macrobiotus hufelandi, Minibiotus furcatus, and Minibiotus intermedius. All have previously been reported from other states and provinces in northeastern North America, but are new records for Maine. Tardigrades (Phylum Tardigrada), commonly known as water bears, are microscopic animals found in marine, freshwater, and terrestrial habitats. Terrestrial species occur in mosses, lichens, liverworts, and leaf litter, and are renowned for their ability to enter a cryptobiotic state (anhydrobiosis) in response to desiccation. Over 900 species of freshwater and terrestrial tardigrade (Phylum Tardigrada) are known worldwide (Degma and Guidetti 2007, Degma et al. 2010, Guidetti and Bertolani 2005). The presence of more than 200 freshwater and terrestrial species has been reported in North America (Meyer and Hinton 2007). Northeastern North America can be defined as a region “roughly bounded from Maryland to Missouri, north to Minnesota and Nunavut, southeast to Newfoundland, and south to West Virginia” (Northeastern Naturalist’s Instructions for Authors). Forty-four published papers and one dissertation have recorded the presence of 99 tardigrade species from 31 genera (Appendix 1) in this region. Although most studies of northeastern North American tardigrades have had a narrow geographical focus, concentrating on a single site or at most a small area, several states and provinces (Illinois, Michigan, New Brunswick, Newfoundland and Labrador, and Virginia) have been more widely surveyed. Records of tardigrades from some states are very limited, with only one or two species identified (e.g., Connecticut and Massachusetts). There are no published records of water bears from Manitoba, Nova Scotia, Prince Edward Island, Delaware, New Jersey, or Rhode Island. The most recent listing of tardigrades from northeastern North America (McInnes 1994) contained no records after the early 1990s, and did not discuss substrates. Northeastern North American tardigrades have mostly been collected from lichens and mosses, with a few records from liverworts, freshwater habitats, leaf litter, and soil (Appendix 1). Not all papers identified the substrate sampled. Many of the papers on the Tardigrada of northeastern North America were written before the development of rigorous standards of specific diagnosis (Guidetti and Bertolani 2005, Pilato and Binda 2001). Some species regarded in older literature as cosmopolitan have proven to belong to complexes of morphologically similar species. Such records must be considered tentative unless confirmed. Nevertheless, in this paper, all species identifications are taken at face value unless emended by subsequent literature. *Department of Biology and Health Sciences, McNeese State University, Lake Charles, LA 70609; hmeyer@mcneese.edu. Notes of the Northeastern Nat u ral ist, Issue 18/4, 2011 2011 Northeastern Naturalist Notes 535 On 24 June 2010, lichen from trees (three samples; mixed foliose, fruticose, and squamulose), moss from rocks (six samples), and deciduous leaf litter (six samples) were collected from the woods near the Bar Harbor Motel, 100 Eden Street, Bar Harbor, Hancock County, ME (44°23'44.9"N, 68°13'33.6"W, 26 m asl). Samples were stored in sealed paper envelopes or paper bags. In the laboratory, samples were soaked in water overnight, thus rehydrating anhydrobiotic water bears. After the samples were sieved (42-μm mesh diameter), they were examined with a dissecting microscope. Tardigrade specimens and eggs were mounted on slides in polyvinyl lactophenol. Sample processing was completed by early October 2010. Tardigrades were identified using keys in Bertolani and Rebecchi (1993), Claxton (1998), Kaczmarek t al. (2011), Nelson and McInnes (2002), Plato and Binda (2010) and Ramazzotti and Maucci (1983), by reference to original descriptions, and by comparison with voucher specimens from elsewhere in North America. No tardigrades were found in the moss samples. Lichen and leaf litter samples contained 139 specimens and 41 eggs, representing seven genera and eleven species (Appendix 1). Echiniscus merokensis, Milnesium tardigradum, Ramazzottius oberhaeuseri, Diphascon (Diphascon) nodulosum, Astatumen trinacriae, Macrobiotus harmsworthi, Macrobiotus hufelandi, Minibiotus furcatus, and Minibiotus intermedius were present in lichen. Leaf litter samples contained Milnesium tardigradum, R. baumanni, R. oberhaeuseri, D. (Adropion) prorsirostre, Macrobiotus harmsworthi, Macrobiotus hufelandi, and Minibiotus intermedius. The presence of eggs of Macrobiotus harmsworthi, Macrobiotus hufelandi, Minibiotus furcatus, and Minibiotus intermedius allowed positive identification of these species. All specimens are deposited in the W.A.K. Seale Museum, McNeese State University, Lake Charles, LA (slides SMLA9700–9720). Packard (1873) observed specimens of tardigrades from water in which moss was growing in New Gloucester, ME, and gave them the invalid name “Macrobiotus americanus”, which cannot be positively identified from his description and drawing. Mathews (1938) mistakenly cited this record as Thulinius augusti, but his identification has not been confirmed. This is the only previous record of water bears from Maine. The eleven species found in this collection therefore constitute new records for the state. All have been previously reported from northeastern North America. Given the small number and spatial extent of the sampling in this preliminary study, doubtless many more tardigrade species remain to be found in Maine. Acknowledgments. I wish to thank Harry L. Meyer and Kathleen R. Jones, who collected the samples. Juliana G. Hinton helped with sample processing. Literature Cited Argue, C.W. 1971. Some terrestrial tardigrades from New Brunswick, Canada. Canadian Journal of Zoology 49:401–415. Argue, C.W. 1972. Tardigrades from New Brunswick, Canada. 2. Canadian Journal of Zoology 50:87–94. Argue, C.W. 1974. Tardigrades from New Brunswick, Canada. 3. Canadian Journal of Zoology 82:919–922. Barrett, G.W., and R.G. Kimmel. 1972. Effects of DDT on the density and diversity of tardigrades. Proceedings of the Iowa Academy of Science 78:41–42. Bateman, L., and M. Collins. 2001. A preliminary account of the tardigrades of Newfoundland. Zoologischer Anzeiger 240:223–232. Beasley, C.W. 1968. The tardigrades of Oklahoma, with additional records from other states and Mexico. Ph.D. Dissertation. University of Oklahoma, Norman, OK. 135 pp. Bernard, E.C. 1977. A new species of Hexapodibius from North America, with a redescription of Diphascon belgicae (Tardigrada). Transactions of the American Microscopical Society 96:476–482. 536 Northeastern Naturalist Vol. 18, No. 4 Bertolani, R., and L. Rebecchi. 1993. A revision of the Macrobiotus hufelandi group (Tardigrada, Macrobiotidae), with some observations on the taxonomic characters of eutardigrades. Zoological Scripta 22:127–152. Boeckner, M., M. Collins, J. Finney-Crawley, and L. Bateman. 2006. The bryofauna of remote coastal Labrador, including a review of current Canadian records. Zootaxa 1105:1–16. Boudrye, M.R. 1967. Notes on the Tardigrada of Minnesota. Proceedings of the Minnesota Academy of Science 26:195–199. Claxton, S. 1998. A revision of the genus Minibiotus (Tardigrada: Macrobiotidae) with description of eleven new species from Australia. Records of the Australian Museum 50:125–160. Curtin, C.B. 1948. The tardigrade fauna of the District of Columbia. Journal of the Washington Academy of Sciences 38:251–254. Curtin, C.B. 1957. Studies on the tardigrades. II. Some tardigrades from Maryland. Pennsylvania Academy of Science 31:142–146. Degma, P., and R. Guidetti. 2007. Notes to the current checklist of Tardigrada. Zootaxa 1579:41–53. Degma P., R. Bertolani, and R. Guidetti. 2010. Actual checklist of Tardigrada species (2009–2011, Ver. 19: 35-05-2011). Available online at http://www.tardigrada.modena.unimo.it/miscellanea/ Actual checklist of Tardigrada.pdf. Accessed 10 June 2011. Evans, W.A. 1982. Abundances of micrometazoans in three sandy beaches in the island area of western Lake Erie. Ohio Journal of Science 82:246–251. Fleeger, J.W., and W.D. Hummon. 1975. Distribution and abundance of soil Tardigrada in cultivated and uncultivated plots of an old field pasture. Memorie dell'Istituto Italiano di Idrobiologia 32 (Suppl.):93–112. Guidetti, R., and R. Bertolani. 2005. Tardigrade taxonomy: An updated checklist of the taxa and a list of characters for their identification. Zootaxa 845:1–46. Hidalgo, H., and D. Coombs. 1985. Tardigrada from Missouri. Transactions of the Kansas Academy of Science 88:121–134. Hohl, A.M., W.R. Miller, and D.R. Nelson. 2001. The distribution of tardigrades upwind and downwind of a Missouri coal-burning power plant. Zoologischer Anzeiger 240:395–401. Iharos, G. 1973. Angaben zur geographischen Verbreitung der Tardigraden. Opuscula Zoologica Budapest 12:73–86. Kaczmarek, Ł., B. Gołdyn, Z. Prokop, and, Ł. Michalczyk. 2011. New records of Tardigrada from Bulgaria with the description of Macrobiotus binieki spec. nov. (Eutardigrada: Macrobiotidae) and a key to the species of the harmsworthi group. Zootaxa 2781:29–29. Kimmel, R.G., and P.A. Meglitsch. 1969. Notes on Iowa tardigrades. Proceedings of the Iowa Academy of Science 76:454–462. Lehmann, R.E., S.D. Shively, and W.R. Miller. 2007. Tardigrades of North America: An historical collection from Kansas and Missouri. Transactions of the Kansas Academy of Science 110:169–178. Mathews, G.B. 1938. Tardigrada from North America. American Midland Naturalist 19:619–627. Maucci, 1987. A contribution to the knowledge of the North American Tardigrada with emphasis on the fauna of Yellowstone National Park (Wyoming). Pp. 187–210, In R. Bertolani (Ed.). Biology of Tardigrades. Selected Symposia and Monographs U.Z.I., 1, Mucchi Editore, Modena, Italy. McInnes, S.J. 1994. Zoogeographic distribution of freshwater/terrestrial tardigrades from the current literature. Journal of Natural History 28:257–352. Mehlen, R.H. 1969. New Tardigrada from Texas. American Midland Naturalist 81:395–404. Meininger, C.A., G.W. Uetz, and J.A. Snider. 1985. Variation in epiphytic microcommunities (tardigrade- lichen-bryophyte assemblages) of the Cincinnati, Ohio area. Urban Ecology 9:45–61. Meyer, H.A., and J.G. Hinton. 2007. Limno-terrestrial Tardigrada of the Nearctic Realm. Journal of Limnology 66 (Suppl. 1):97–103. Meyer, H.A., A.M. Lyons, D.R. Nelson, and J.G. Hinton. 2011. Terrestrial Tardigrada of Michigan, northern USA, with the description of Minibiotus jonesorum sp. n. (Eutardigrada: Macrobiotidae). Journal of Systematic Zoology and Evolutionary Research 4 (Suppl. 1):1–8. Mitchell, C.R., W.R. Miller, and B. Davis. 2009. Tardigrades of North America: Influence of substrate on habitat selection. Journal of the Pennsylvania Academy of Science 83:10–16 Murray, J. 1910. Tardigrada. Report on Scientific Investigations of the British Antarctic Expedition 1907–1909 1:81–185. 2011 Northeastern Naturalist Notes 537 Nelson, D.R. 1991. A new species of Diphascon from New Brunswick, Canada (Tardigrada). Canadian Journal of Zoology 69:1911–1915. Nelson, D.R., and S.J. McInnes. 2002. Tardigrada. Pp. 177–215, In S.D. Rundle, A. Robertson, and J.M. Schmid-Araya (Eds.). Freshwater Meiofauna: Biology and Ecology. Backhuys Publishers, Leiden, Netherlands. 370 pp. Packard, A.S. 1873. Discovery of a tardigrade. American Naturalist 7:740–741. Pilato, G., and M.G. Binda. 2001. Biogeography and limno-terrestrial tardigrades: Are they truly incompatible binomials? Zoologischer Anzeiger 240:511–516. Pilato, G., and M.G. Binda. 2010. Definition of families, subfamilies, genera and subgenera of the Eutardigrada, and keys to their identification. Zootaxa 2404:1–54. Puglia, C.H. 1964. Some tardigrades from Illinois. Transactions of the American Microscopical Society 83:300–311. Ramazzotti, G. 1957. Due nueve specie di tardigrade extraeuropei. Atti della Società Italiana di Scienze Naturali e del Museo Civico di Storia Naturale di Milano 96:188–191. Ramazzotti, G., and W. Maucci. 1983. Il Phylum Tardigrada. Memorie dell’Istituto Italiano di Idrobiologia 41:1–1011. Riggin, G.T., Jr. 1962. The Tardigrada of southwest Virginia: With the addition of the description of a new marine species from Florida. Virginia Agricultural Experiment Station, Technical Bulletin 152:1–145. Riggin, G.T., Jr. 1964. Tardigrada from the southern Appalachian Mountains. Transactions of the American Microscopical Society 83:277–282. Ryan, J. 1977. Invertebrates of Truelove Lowland. Pp. 699–703, In L.C. Bliss (Ed.). Truelove Lowland, Devon Island, Canada: A High Arctic Ecosystem. University of Alberta Press, Edmonton, AB. 714 pp. Strayer, D. 1985. The benthic micrometazoans of Mirror Lake, New Hampshire. Archiv für Hydrobiologie 72 (Suppl.):287–426. Strayer, D., D.R. Nelson, and E.B. O’Donnell. 1994. Tardigrades from shallow groundwaters in southeastern New York, with the first record of Thulinia from North America. Transactions of the American Microscopical Society 113:325–332. Szymanska, B. 1994. The Tardigrada from the Axel Heiberg Island and the associated bryophyte species. Folia Entomologica Hungarica 55:359–368. Tarter, D., and D. Nelson. 1993. First records of tardigrades (Phylum: Tardigrada) from mosses in the Cranberry Glades Botanical Area in the Monongahela National Forest, Pocahontas County, West Virginia. Proceedings of the West Virginia Academy of Science 65:36. Tarter, D., and D. Nelson. 1994. Preliminary list of tardigrades (Phylum: Tardigrada) from mosses and liverworts in the Monongahela National Forest. Proceedings of the West Virginia Academy of Science 66:34. Tarter, D., K. Ruggles, S. Gillenwater, and D. Nelson. 1992. First records of water bears (Phylum: Tardigrada) from West Virginia. Proceedings of the West Virginia Academy of Science, 1989 61:96–99. Van Rompu, E.A., W.H. De Smet, and J.M. Bafort. 1991. Contributions to the Tardigrada of the Canadian High Arctic. 2. Freshwater tardigrades from the Little Cornwallis Island, Northwest Territories, Canada. Biologisch Jaarboek Dodonaea 59:132–140. Van Rompu, E.A., W.H. De Smet, and L. Beyens. 1992. Contributions to the Tardigrada of the Canadian High-Arctic 1. Freshwater tardigrades from Devon Island, Northwest Territories. Canadian Field-Naturalist 106:303–310. Van Rompu, E.A., W.H. de Smet, and L. Beyens. 1999 (2000). Tardigrada from Victoria Island, Arctic Canada. Biologisch Jaarboek Dodonaea, 67:124–137. Wainberg, R.H., and W.D. Hummon. 1981. Morphological variation of the tardigrade Isohypsibius saltursus. Transactions of the American Microscopical Society 100:21–33. Węglarska, B. 1970. Hypsibius (Isohypsibius) smreczynskii spec. nov., a new species of freshwater tardigrade. Zeszyty Naukowe Uniwersytetu Jagiellonskiego Prace Zoologicne 16:107–114. Węglarska, B., and M. Kuc. 1980. Heterotardigrada from Axel Heiberg Island. Zeszyty Naukowe Uniwersytetu Jagiellonskiego Prace Zoologicne 26:53–66. Winkelmann, R.K., C. Elzinga, D.R. Nelson, M.B. Edlund, and M.R. Sommerfeld. 2004. The plankton of the St. Croix River in 2000, including tardigrades, a red alga, and Chrysophyceae. Journal of Freshwater Ecology 19:179–188. 538 Northeastern Naturalist Vol. 18, No. 4 Appendix 1. Terrestrial and freshwater tardigrades of northeastern North America (excluding Greenland). Locations: DC = District of Columbia; Canadian provinces – NB = New Brunswick, NF = Newfoundland and Labrador, NU = Nunavut, ON = Ontario, QC = Quebec ; USA states – Connecticut = CT, IL = Illinois, IO = Iowa, MA = Massachusetts, MD = Maryland, ME = Maine, MI = Michigan, MN = Minnesota, MO = Missouri, NH = New Hampshire, NY = New York, OH = Ohio, PA = Pennsylvania, VA = Virginia, Vermont = VT, WV = West Virginia, and WI = Wisconsin. Citations: A1 = Argue (1971), A2 = Argue (1972), A3 = Argue (1974), BC = Bateman and Collins (2001), BCF = Boeckner et al. (2006), Bea = Beasley (1968), Ber = Bernard (1977), BK = Barrett and Kimmel (1972), Bo = Boudrye (1967), C1 = Curtin (1948), C2 = Curtin (1957), E = Evans (1982), FH = Fleeger and Hummon (1975), HC = Hidalgo and Coombs (1985), HW = Hohl et al. (2001), I = Iharos (1973), K = Kimmel and Meglitsch (1969), L = Lehmann et al. (2007), Mat = Mathews (1938), Mau = Maucci (1987), Meh = Mehlen (1969), Mey = this paper, MH = Meyer and Hinton (2007), ML = Meyer et al. (2011), MM = Mitchell et al. (2009), MU = Meininger et al. (1985), Mu = Murray (1910), N = Nelson (1991), P = Puglia (1964), Ra = Ramazzotti (1957), Ri1 = Riggin (1962), Ri2 = Riggin (1964), Ry = Ryan (1977), St = Strayer (1985), SN = Strayer et al. (1994), Sz = Szymanska (1994), TN1 = Tarter and Nelson (1993), TN2 = Tarter and Nelson (1994), TR = Tarter et al. (1992), V1 = Van Rompu et al. (1991), V2 = Van Rompu et al. (1992), V3 = Van Rompu et al. (1999 [2000]), W = Węglarska (1970), WE = Winkelmann et al. (2004), WH = Wainberg and Hummon (1981), and WK = Węglarska and Kuc (1980). Substrates: C = Cryptogam (unspecified lichen, liverwort, or moss), Fw = Benthic freshwater, Gw = groundwater, Lc = Lichen, Ll = Leaf litter, Lv = Liverwort, M = Moss, S = Soil, ? = substrate not identified. Species Location (Citations) Substrate Bryodelphax parvulus Thulin, 1928 NU (V3) Fw Cornechiniscus cornutus (Richters, 1906) NU (Sz, WK) M Cornechiniscus holmeni (Petersen, 1951) NU (Sz, WK) M Echiniscus sp. MN (Bo), MO (HW) Lc Echiniscus arctomys Ehrenberg, 1853 VT (Mat), WI (Mat) ? Echiniscus blumi Richters, 1903 MI (ML), NU (Sz, WK) M Echiniscus dearmatus Bartoš, 1935 QC (I) M Echiniscus kerguelensis Richters, 1904 NU (Sz, WK) M Echiniscus mauccii Ramazzotti, 1956 MI (ML), MO (HC), NB (A1), Lc OH (MU) Echiniscus merokensis Richters, 1904 ME (Mey), MI (ML), NF (BC), Lc, M NU (Sz, WK) Echiniscus quadrispinosus Richters, 1902 NU (Sz, WK) M Echiniscus spiniger Richters, 1904 NB (A2) Lc, Lv, M Echiniscus sylvanus Murray, 1910 ON (Mat, Mu) M Echiniscus testudo (Doyère, 1840) NU (Sz, WK) M Echiniscus trisetosus Cuénot, 1932 NU (Sz, WK) M Echiniscus virginicus Riggin, 1962 VA (Ri1), MI (ML) Lc, M Echiniscus wendti Richters, 1903 IL (P), MI (ML), MO (L), Lc, M NF (BC, BCF), NU (Sz, WK) Hypechiniscus gladiator (Murray, 1905) NB (A1), VA (Ri1), WV (TN2, TR) M Proechiniscus hanneae (Petersen, 1951) NF (BC, BCF) M Pseudechiniscus sp. NU (Ry) ? Pseudechiniscus facettalis Petersen, 1951 NU (WK) M Pseudechiniscus suillus (Ehrenberg, 1853) MI (ML), NB (A1), NU (Sz, V2,3, Fw, M WK), VA (Ri1), VT (Mat), WV (TN2) Pseudechiniscus victor (Ehrenberg, 1853) NU (Sz, V3, WK) Fw, M Testechiniscus spitsbergensis (Scourfield, 1897) NB (A2), NU (Sz, V1–3, WK) Fw, M Milnesium sp. MN (Bo) Lc 2011 Northeastern Naturalist Notes 539 Species Location (Citations) Substrate Milnesium tardigradum Doyère, 1840 DC (C1), IL (Mat), IL (P), Lc, Ll, M IO (BK, K), MD (C2), ME (Mey), MI (ML), MO (HC, HW, L), NB (A1), NF (BC, BCF), NU (Sz), OH (MU), ON (Mat, Mu), PA (MM),QC (I), VA (Ri1), VT (MH),WI (Mat, Mau), WV (Bea, TN1, TR) Bertolanius smreczynskii (Węglarska, 1970) NU (Sz, W) Fw Bertolanius weglarskae (Dastych, 1972) NU (V2, V3) Fw Calohypsibius ornatus (Richters, 1900) NF (BC, BCF), NU (Sz) M Parhexapodibius pilatoi (Bernard, 1977) MI (Ber) S Borealibius zetlandicus (Murray, 1907) NU (Sz) M Hypsibius sp. DC (C1) Fw Hypsibius arcticus (Murray, 1907) MI (ML), ON (Mu) M Hypsibius calcaratus Bartoš, 1935 MI (ML) Ll Hypsibius convergens (Urbanowicz, 1925) MD (C2), MO (L), NB (A1), NF Fw, Lc, M (BC, BCF), NU (Sz, V1–3), NY (Mat), ON (Mat), VA (Ri1), WV (TN1, TR) Hypsibius dujardini (Doyère, 1840) DC (C1), MD (C2), MO (L), NB Fw, M, S (A3), NF (BC, BCF), NU (Sz, V1,2), OH (FH), WV (TR) Hypsibius maculatus Iharos, 1969 WV (TN2, TR) C Hypsibius microps Thulin, 1928 NF (BC, BCF) M Hypsibius pallidus Thulin, 1911 NB (A1), NF (BC), NU (Sz, V3), Fw, M QC (I) Isohypsibius sp. MN (WE), NU (V3), NY (SN) Fw, Gw Isohypsibius cf. baldii NY (SN) Gw Isohypsibius canadensis (Murray, 1910) VA (Ri1) C Isohypsibius cf. canadensis NU (V2) Fw Isohypsibius elegans Binda and Pilato, 1971 NU (V3) Fw Isohypsibius granulifer Thulin, 1928 NU (V1–3) Fw Isohypsibius lunulatus (Iharos, 1966) NF (BC) M Isohypsibius macrodactylus (Maucci, 1978) WV (TR) M Isohypsibius nodosus (Murray, 1907) IL (P), NU (Sz), VA (Ri1) Ll, M Isohypsibius papillifer (Murray, 1905) NU (V1–3) Fw Isohypsibius prosostomus Thulin, 1928 MI (Mat), NF (BC), OH (FH), M, S VA (Ri1) Isohypsibius sattleri (Richters, 1902) NB (A1), NU (Sz), WV (TN1) M Isohypsibius schaudinni (Richters, 1909) MI (ML), NF (BC), NU (Sz, V2), Fw, Lc, QC (I) Ll, M Isohypsibius tetradactyloides (Richters, 1907) NU (Sz, V1,2), VA (Ri1) Fw, M Isohypsibius tuberculatus (Plate, 1888) IO (K), NB (A1), OH (FH), Lc. M, S VA (Ri1) Pseudobiotus sp. NH (St) Fw Ramazzottius sp. MO (HW), NF (BC) Lc, M Ramazzottius baumanni (Ramazzotti, 1962) ME (Mey), MI (ML) Lc, Ll, M Ramazzottius oberhaeuseri (Doyère, 1840) IL (P), ME (Mey), MI (ML), Lc, Ll, M NB (A1), NU (Sz), PA (MM), QC (I), VA (Ri1), WI (Mat), WV (TN1,2, TR) Thulinius augusti (Murray, 1907) IL (P), OH (WH), ON (E) Fw 540 Northeastern Naturalist Vol. 18, No. 4 Species Location (Citations) Substrate Thulinius ruffoi (R. Bertolani, 1982) NY (SN) Gw Thulinius saltursus (Schuster, Toftner, OH (WH), ON (E) Fw and Grigarick, 1978) Thulinius stephaniae (Pilato, 1974) NU (V3) Fw Diphascon sp. WV (TR) M Diphascon (Diphascon) alpinum Murray, 1906 MI (ML), ON (Mu) Lc, Ll, M Diphascon (Diphascon) brevipes (Marcus, 1936) IL (P), MO (L), NU (Sz) Ll, M, So Diphascon (Diphascon) bullatum Murray, 1905 NB (A2), OH (FH), VA (Ri1) M, S Diphascon (Diphascon) burti Nelson, 1991 NB (N) C Diphascon (Diphascon) chilenense Plate, 1888 NU (Sz) M Diphascon (Diphascon) nodulosum ME (Mey), MI (ML), NB (A2), Lc, Ll, M (Ramazzotti, 1957) NF (BC, BCF), VA (Ri1), WI (Ra) Diphascon (Diphascon) oculatum Murray, 1906 NU (Sz) M Diphascon (Diphascon) ongulense NU (V3) Fw (Morikawa, 1962) Diphascon (Diphascon) pingue (Marcus, 1936) CT (Ri1), NB (A2), NF (BC, BCF), Fw, M NU (Sz, V3), VA (Ri1), WI (Mau) Diphascon (Diphascon) ramazzottii NF (BC, BCF) M (Robotti, 1971) Diphascon (Diphascon) recamieri Richters, 1911 NF (BCF), NU (Sz, V2) Fw. M Diphascon (Diphascon) rugosum Bartoš, 1935 NB (A3), NU (V3) Fw, M Diphascon (Diphascon) tenue Thulin, 1928 QC (I) M Diphascon (Adropion) belgicae Richters, 1911 MI (Ber), MO (L), NB (A1), M, S OH (FH), VA (Ri1) Diphascon (Adropion) prorsirostre Thulin, 1928 ME (Mey), MI (ML), NB (A1), M, Ll WV (TN2, TR) Diphascon (Adropion) scoticum Murray, 1905 MI (ML), MO (L), NB (A1), Ll, M NF (BC, BCF), OH (MU), VA (Ri1), WV (TN2, TR) Hebesuncus conjungens (Thulin, 1911) MD (C2), NF (BC), NU (Sz), Lc, M VA (Ri1) Astatumen bartosi (Węglarska, 1959) OH (MU) C Astatumen trinacriae (Arcidiacono, 1962) ME (Mey), MI (ML) Ll, M Itaquason sp. PA (MM) Lc, M Itaquascon placophorum Maucci, 1973 WV (TR) M Itaquascon umbellinae de Barros, 1939 IL (P), VA (Ri1) Lc, Ll, M Mesocrista spitsbergensis (Richters, 1903) NB (A1), NF (BC, BCF) M Platicrista angustata (Murray, 1905) MO (L), NB (A1), NF (BC, BCF), M NU (Sz), VA (Ri1), VT (Ri2) Macrobiotus sp. MN (Bo), MO (HW), NH (St), Fw, Lc NU (Ry) Macrobiotus crenulatus Richters, 1904 NF (BCF) M Macrobiotus echinogenitus Richters, 1904 MD (C2), MI (Bea, ML), Lc, M MO (HC, L), NB (A1), NU (Sz), ON (Mu), VA (Ri1) Macrobiotus furciger Murray, 1907 NB (A2) M Macrobiotus harmsworthi Murray, 1907 IL (P), ME (Mey), MI (ML), Fw, Lc, MO (HC, L), NB (A1), NF (BC), Ll, M NU (Sz, V1), ON (Mat, Mu), PA (MM), VA (Ri1), VT (Ri2), WV (TN1, TR) Macrobiotus cf. harmsworthi NF (BC, BCF) M Macrobiotus hibiscus de Barros, 1942 OH (MU) C 2011 Northeastern Naturalist Notes 541 Species Location (Citations) Substrate Macrobiotus hufelandi C.A.S. Schultze, 1833 DC (C1), IL (P), IO (KM), Lc, Ll, M, ME (Mey), MI (Mat, ML), S MO (HC, L), NB (A1), NF (BC), NU (Sz), OH (FH, MU), PA (MM), QC (I), VA (Ri1), VT (Ri2), WI (Mat), WV (TR) Macrobiotus cf. hufelandi NF (BC, BCF) M Macrobiotus islandicus Richters, 1904 IO (BK, KM), PA (MM) Lc, M Macrobiotus montanus Murray, 1910 NB (A1), NF (BC), NU (Sz) M Macrobiotus occidentalis Murray, 1910 MA (Meh), NU (Sz), M WV (TN2, TR) Macrobiotus pseudofurcatus Pilato, 1972 WV (TN1) M Macrobiotus spectabilis Thulin, 1928 WV (TN1) M Macrobiotus virgatus Murray, 1910 ON (Mu) C Minibiotus sp. MO (HW) Lc Minibiotus furcatus (Ehrenberg, 1859) IL (P), ME (Mey), MO (L), Lc, Ll VT (MH) Minibiotus hufelandioides (Murray, 1910) MI (ML) Lc Minibiotus intermedius (Plate, 1888) IL (P), MD (C2), MA (Meh), Lc, Ll, M ME (Mey), MI (ML), MO (HC, L), NB (A1), NF (BC, BCF), NU (Sz), OH (MU),ON (Mat, Mu), PA (MM),VA (Ri1), WI (Mau), WV (TN1, TR) Minibiotus jonesorum Meyer, Lyons, Nelson MI (ML) Lc and Hinton, 2011 Paramacrobiotus areolatus (Murray, 1907) IO (BK, KM), MI (ML), Lc, Ll, M MO (HC, L), NU (Sz), ON (Mu), PA (MM), WV (Bea) Paramacrobiotus richtersi (Murray, 1911) IL (P), MI (ML), NB (A1), Ll, M NF (BC),NU (Sz), QC (I), VA (Ri1), VT (Ri2), WV (TR) Paramacrobiotus tonollii (Ramazzotti, 1956) IL (P), MI (ML), MO (HC, L), Lc, Ll, M OH (MU), VT (MH), WI (Mau), WV (TR) Richtersius coronifer (Richters, 1903) NU (Sz) M Dactylobiotus dispar (Murray, 1907) MO (L), NU (V1–3), VA (Ri2) Fw Dactylobiotus cf. dispar NY (SN) Gw Dactylobiotus grandipes (Schuster, Toftner NH (St) Fw and Grigarick, 1978) Dactylobiotus ambiguus (Murray, 1907) NB (A2), NU (Sz) M Dactylobiotus macronyx (Dujardin, 1851) VA (Ri1) Fw Murrayon dianeae (Kristensen, 1982) NF (BC), NU (V1–3) Fw, M Murrayon hibernicus (Murray, 1911) NB (A2) M Murrayon pullari (Murray, 1907) NB (A3) Fw Murrayon pullari/nocentiniae NY (SN) Gw Fractonotus caelatus (Marcus, 1928) MI (ML) M Microhypsibius sp. NF (BC) M