2008 SOUTHEASTERN NATURALIST 7(1):91–100 Distribution of Tardigrades in Florida Harry A. Meyer* Abstract - The distribution of terrestrial tardigrades in the Gulf Coast states of the United States is poorly known. Only one species has been reported in Florida. In this study, bryophyte samples (47 identified species) from trees and shrubs were collected from all 67 Florida counties. Twenty tardigrade species were found: Echiniscus cavagnaroi, E. kofordi, E. virginicus, Pseudechiniscus suillus, Ramazzottius baumanni, Diphascon (Diphascon) pingue, Macrobiotus echinogenitus, two species of Macrobiotus cf. harmsworthi, Macrobiotus cf. hibiscus, Macrobiotus hufelandi, two species of Macrobiotus cf. hufelandi, Macrobiotus islandicus, Macrobiotus occidentalis, Macrobiotus richtersi, Macrobiotus tonollii, Minibiotus fallax, Minibiotus intermedius, and Milnesium tardigradum. The distribution of tardigrade species was not correlated with Florida’s ecological regions. Tardigrade species richness declined from north to south in peninsular Florida. Introduction Until recently, the distribution of terrestrial tardigrades in the states of the Gulf Coast of the United States of America (Texas, Louisiana, Mississippi, Alabama, and Florida) was virtually unknown. Only in Texas had extensive sampling been carried out. Three studies (Caskey 1971, Mathews 1938, Mehlen 1969) reported a total of 18 species in that state. Tardigrades were unknown in Louisiana and Mississippi, and only four species were known in Alabama and Florida (Christenberry 1979, Christenberry and Higgins 1979, Christenberry and Mason 1979). More recent studies have added considerably to our knowledge of the tardigrade fauna of Louisiana (Meyer 2001) and Alabama (Nichols et al. 2001, Romano et al. 2001). Although Meyer (2006a) published a list of Florida tardigrades, no distribution data were included. The state of Florida consists of the peninsula and adjacent mainland counties, and a western extension (the panhandle) along the Gulf Coast. No point in Florida is more than 100 km from the Gulf of Mexico or the Atlantic Ocean. The maximum elevation is 120 m above sea level. Florida spans 6.5 degrees of latitude, from a temperate continental north to an essentially tropical south. The incidence of frost declines with decreasing latitude, and winter freezes rarely occur in the southern third of the peninsula (Chen and Gerber 1990). Although the Florida peninsula is about 25 million years old, its area has fl uctuated widely with changes in sea level (Webb 1990). While the mesic forests and xeric pine barrens, sandhills, and scrublands of the interior have existed for 20–25 million years, coastal and lakeshore habitats are only a few thousand years old (Webb 1990). *Department of Biological and Environmental Sciences, McNeese State University, Lake Charles, LA; hmeyer@mcneese.edu. 92 Southeastern Naturalist Vol.7, No. 1 The United States Department of Agriculture (USDA) Forest Service classifies Florida into four ecological sections (Fig. 1). All four sections are within the Humid Temperate Domain. The Subtropical Division includes the Lower Coastal Plains and Flatwoods Section (most mainland and interior counties), and the Eastern and Western Florida Coastal Lowlands Sections. The Everglades, in the southern third of the peninsula, constitute the fourth section, assigned to the Savanna Division. The geography of peninsulas, like islands, may affect species diversity. Simpson (1964) proposed a “peninsular effect,” which predicts that species richness will decline along a peninsula from its base to its tip, and that a peninsula will have fewer species than equivalent areas of mainland. The prediction assumes that colonization from the mainland is the source of peninsular species. Studies assessing the validity of Simpson (1964)’s prediction in various peninsulas have yielded inconsistent results. Mears and Simberloff (1987) found the predicted decline in the diversity of amphibians and reptiles in peninsular Florida, and Choi (2004) showed that butterfl y diversity declines from north to south in Korea. Johnson and Ward (2002), on the other hand, found little support for the predicted decline in ant diversity in Baja California, where habitat diversity had a much stronger effect on species richness . I collected lichens, mosses (phylum Bryophyta), liverworts (phylum Hepatophyta), and Pleopeltis polypodioides (L.) Andrews and Windham (resurrection fern) from trees, bushes, and fallen branches at sites throughout Florida. These potential tardigrade substrates will hereafter be collectively referred to as cryptogams. The data were examined for evidence of correlation of tardigrade distribution patterns with the Forest Service’s ecological sections, and Simpson’s peninsular effect. Material and Methods Tardigrades were collected from 1999 to 2003 from 83 sites encompassing all 67 counties of Florida (Fig. 1). The Florida Keys and other offshore islands were not visited. Collecting season varied among the sites. At each site, suitable tardigrade substrates were collected from epiphytic cryptogams on trees, shrubs, and fallen branches. Cryptogam samples were approximately the same size (handfuls). Exposed rocks are relatively rare in Florida, and material from only one such habitat (Site 10: a limestone outcrop) was examined. The latitude, longitude, and altitude of each site were recorded. Where possible, sites were chosen in national and state parks and forests. However, because counties often lack such sites, many samples were collected along rural roads and even in highly disturbed urban habitats. Supplementary Table 1 (available only online at http://dx.doi.org/10.1656/S407.s1) lists and describes all sites, which will hereafter be referred to by their number (including an additional letter if more than one cryptogam was collected at the site). In most cases, tree and shrub species were not identified. Where possible, cryptogams were 2008 H.A. Meyer 93 identified to species, using keys and illustrations provided in Breen (1963), Bordo et al. (2001), Conard (1956), Hale (1969), and Reese (1983). Handfuls of substrate were stored in sealed paper envelopes and allowed to dry slowly. An entire sample was placed in deionized water, and after soaking for 1–3 days, the material was sorted using a dissecting microscope and mounted in modified Hoyer’s medium (Christenberry 1979). After drying in a 65 oC oven for several hours (Hohl et al. 2001), the slides were ringed with clear fingernail polish. Morphological measurements were made with an ocular micrometer. Tardigrades were identified using keys and descriptions in Bertolani and Rebecchi (1993), Nelson and McInnes (2002), and Ramazzotti and Maucci (1983), and by reference to the primary literature. The data were inspected visually for a relationship between tardigrade distribution and the four USDA ecological regions. Figure 1. Map of Florida showing collecting sites and division of the state into three north-south regions (solid lines) and four ecological regions (dotted lines). 232B = Lower Coastal Plains and Flatwood Section, 232D = Florida Coastal Lowlands (Western) Section, 232G = Florida Coastal Lowlands (Eastern) Section, 411A = Everglades. 94 Southeastern Naturalist Vol.7, No. 1 Simpson (1964)’s prediction that species richness in a peninsula declines from base to tip was investigated by dividing peninsular Florida (i.e., excluding the panhandle and the northern counties along the Georgia border) into three regions from north to south (Fig. 1). Sites were assigned to regions so as to equalize, as much as possible, the number of samples per region. Although this division is arbitrary, the boundary between the middle and southern regions corresponds to the northern edge of the Everglades ecosystem. Simpson (1964)’s prediction that species richness in a peninsula is lower than for equivalent mainland areas could not be evaluated, since almost nothing is known of terrestrial tardigrades in Georgia. Results Cryptogams Tardigrades were collected from 47 identified species of cryptogam (Supplementary Table 1; available only online at http://dx.doi.org/ 10.1656/S407.s1). These cryptogam species included 13 mosses, 3 liverworts, and 31 lichens. Some additional cryptogam samples were not identified beyond genus or general category (e.g., moss, foliose lichen, etc.). No tardigrades were found in samples of five cryptogam species: the moss Aulocomnium heterostichum (Hedw.) B.S.G., the lichens Physcia neogaea R.C. Harris, Ramalina dilacerata (Hoffm.) Hoffm., and Usnea mutabilis Stirton, and the fern Pleopeltis polypodioides. Tardigrades A total of 1523 tardigrades and eggs, representing 20 species, was collected from Florida cryptogams (Appendix 1). Four species belong to the class Heterotardigrada and the remainder to the class Eutardigrada. While the maximum number of species found in one sample was four, the modal number was two. Ecological regions and peninsular effect Visual inspection of the data did not reveal any trend relating tardigrade distribution in Florida to the four USDA ecological sections. The more common species were all found in two or more of the sections. Species richness of tardigrades declined from north to south in peninsular Florida (15 species in the northern portion, 11 in the middle, and 9 in the southern region). The number of species per sample also appeared to decline. However, since sampling was not quantitative, this trend could not be tested statistically. Discussion This survey found only 20 species of tardigrades in Florida cryptogams. Given that I was unable to include samples from all sites throughout all 2008 H.A. Meyer 95 seasons, this study should be considered a preliminary survey of tardigrade diversity in Florida. No doubt additional species remain to be found. Most of the tardigrade species in Florida have widespread distributions (McInnes 1994). Milnesium tardigradum, Macrobiotus echinogenitus, Macrobiotus hufelandi, Macrobiotus occidentalis, Macrobiotus richtersi, Minibiotus intermedius, Diphascon (Diphascon) pingue, and Pseudechiniscus suillus all have global distributions. The distribution of Macrobiotus islandicus is Holarctic. Macrobiotus tonollii is found throughout much of North America. Ramazzottius baumanni has been collected throughout the western hemisphere. Echiniscus virginicus, E. kofordi, and E. cavagnaroi are widely distributed in the southern United States (Hinton and Meyer 2007). Minibiotus fallax was previously known only in Australia (Pilato et al. 1989). It also appears to be widely distributed in the southern United States since it is also found in Louisiana (Hinton and Meyer 2007). Among the tardigrade species that could not be attributed to known taxa, Macrobiotus cf. hufelandi sp. 2 has also been found in Louisiana (Hinton and Meyer 2007). The number of tardigrade species found in a cryptogam sample usually ranges from two to six, although more than ten are sometimes found (Ramazzotti and Maucci 1983). Tardigrade species richness in individual cryptogams in Florida, ranging from one to four with a mode of two, is at the lower end of this global average, and is characteristic of tardigrades in lichens from disturbed or xeric habitats (Séméria 1982). Typically, such substrates have one predatory species and several detrivorous or herbivorous species (Séméria 1982). The Florida data suggest a similar pattern for tardigrades in cryptogams. One may envision Florida cryptogams as having four potential niches for tardigrade species: one predatory species (Milnesium tardigradum, Macrobiotus cf. harmsworthi or Macrobiotus richtersi), one other Macrobiotus species, one Minibiotus species, and one Echiniscid species. Although in many cryptogams, one or more of these slots may be unoccupied, in virtually every sample, the occupied slots followed these rules (the only exception was one sample in which two predatory species were present). Fewer tardigrade species were found in samples from the southern portion of peninsular Florida than in those from the north. This decline is explained by the distribution of species uncommon in the samples. Most of these species (those found at only one or two sites) were found in the northern portion of peninsular Florida (e.g., Ramazzottius baumanni, Macrobiotus islandicus, and Macrobiotus occidentalis). All tardigrade species found in the southernmost counties of Florida (e.g., Macrobiotus cf. hufelandi sp. 2, Minibiotus intermedius, and Milnesium tardigradum) were also found widely throughout the state. Available data on the distribution of tardigrades in peninsular Florida appears to support Simpson’s prediction of a peninsular effect. 96 Southeastern Naturalist Vol.7, No. 1 The data do not suggest that tardigrade distribution differs among ecological sections as defined by the Forest Service. The environmental and vegetation differences distinguishing sections may be too coarse-grained to influence tardigrade distribution, which can vary enormously even on very small spatial scales (Meyer 2006b). Another possibility is that human disruption of Florida vegetation is severe enough to obscure differences among sections. Human impact on Florida ecosystems has been enormous and continues to grow (Ewel 1990). Much of the original habitat has been urbanized or converted to farmlands and forestry plantations. These anthropogenic impacts may well affect tardigrade distributions. Séméria (1982) suggested that the tardigrade fauna of such disrupted habitats, responding to the altered biotic and environmental milieu, may have their own characteristic assemblages, which need not be identical to those in “natural” habitats of the same area. This possibility merits further investigation in Florida and elsewhere. Acknowledgments Harry L. Meyer, H. Thomas Meyer, and Daniel H. Jones helped with field collection. M. Ray Neyland identified some plants. Juliana G. Hinton, Remigius J. Jackson, Jwill Sims, and Kathleen Trahan helped with sample processing. This research was supported by the award to the author of a Franklin L. and Laura Chavanne Miller Professorship in Science. Literature Cited Bertolani, R., and L. Rebecchi. 1993. A revision of the Macrobiotus hufelandi group (Tardigrada, Macrobiotidae), with some observations on the taxonomic characters of eutardigrades. Zoologica Scripta 22:127–152. Breen, R.S. 1963. Mosses of Florida: An Illustrated Manual. University of Florida Press, Gainesville, FL. 273 pp. Brodo, I.M., S.D. Sharnoff, and S. Sharnoff. 2001. Lichens of North America. Yale University Press, New Haven, CT. 793 pp. Caskey, D.S. 1971. Tardigrada of Texas. M.Sc. Thesis. Lamar University, Beaumont, TX. 88 pp. Chen, E., and J.F. Gerber. 1990. Climate. Pp. 11–34, In R.L Myers and J.J. Ewel (Eds.). Ecosystems of Florida. University of Central Florida Press, Orlando, FL. Choi, S. 2004. Trends in butterfly richness in response to the peninsular effect in South Korea. Journal of Biogeography 31:587–592. Christenberry, D. 1979. On the distribution of Echiniscus kofordi and E. cavagnaroi (Tardigrada). Transactions of the American Microscopical Society 98: 469–471. Christenberry, D., and R.P. Higgins. 1979. A new species of Pseudodiphascon (Tardigrada) from Alabama. Transactions of the American Microscopical Society 98:508–514. Christenberry, D., and W.H. Mason, 1979. Redescription of Echiniscus virginicus Riggin with notes on life history, range, and geographic variation. Journal of the Alabama Academy of Science 50:47–61. 2008 H.A. Meyer 97 Conard, H.S., Jr. 1956. How to Know the Mosses and Liverworts. Revised Edition. Wm. C. Brown Company Publishers, Dubuque, IA. 226 pp. Ewel, J.J. 1990. Introduction. Pp. 3–10 and 34, In R.L Myers and J.J. Ewel (Eds.). Ecosystems of Florida. University of Central Florida Press, Orlando, FL. Grigarick, A.A., R.O. Schuster, and D.R. Nelson. 1983. Heterotardigrada of Venezuela (Tardigrada). Pan Pacific Entomologist 59:64–77. Hale, M. E. 1969. How to Know the Lichens. Wm. C. Brown Company Publishers, Dubuque, IA. 226 pp. Hinton, J.G., and H.A. Meyer. 2007. Distribution and ecology of tardigrades in the Gulf Coast states of the United States of America. Journal of Limnology 66(Suppl.1):72–76. 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. Johnson, R.A., and P.S. Ward. 2002. Biogeography and endemism of ants (Hymenoptera: Formicidae) in Baja California, Mexico: A first overview. Journal of Biogeography 29:1009–1026. Mathews, G.B. 1938. Tardigrada from North America. American Midland Naturalist 81:395–404. McInnes, S.J. 1994. Zoogeographic distribution of terrestrial/freshwater tardigrades from current literature. Journal of Natural History 28:257–352. Mears, D.B., and D. Simberloff. 1987. The peninsular effect: Habitat-correlated species decline in Florida’s herpetofauna. Journal of Biogeography 14:551–568. Mehlen, R. 1969. New Tardigrada from Texas. American Midland Naturalist 81: 395–404. Meyer, H.A. 2001. Tardigrades of Louisiana and Arkansas, United States of America. Zoologischer Anzeiger 240:471–474. Meyer, H.A. 2006a. Interspecific association and substrate specificity in tardigrades from Florida. Hydrobiologia 558:129–132. Meyer, H.A. 2006b. Small-scale spatial distribution variability in terrestrial tardigrade populations. Hydrobiologia 558:133–139. Nelson, D.R., and S.J. McInnes. 2002. Tardigrada. Pp. 177–215, In S.D. Rundle, A.L.Robertson, and J.M. Schmid-Araya (Eds.). Freshwater Meiofauna: Biology and Ecology. Backhuys Publishers, Leiden, The Netherlands. Nichols, P.B., F.A. Romano III, and D.R. Nelson. 2001. Seasonal and altitudinal variation in the distribution and abundance of Tardigrada on Dugger Mountain, Alabama. Zoologischer Anzeiger 240:501–504. Pilato, G., S. Claxton, and M.G. Binda. 1989. Tardigrades from Australia. II. The evaluation of Calohypsibius ornatus (Richters, 1900) caelatus (Marcus, 1928) as a valid species and description of Minibiotus fallax n. sp. (Eutardigrada). Animalia 16:21–27. Ramazzotti, G., and W. Maucci. 1983. Il Philum Tardigrada. Memorie dell’Istituto Italiano di Idrobiologia 41:1–1011. Reese, W.D. 1983. Mosses of the Gulf South: From the Rio Grande to the Aplachicola. Louisiana State University, Baton Rouge, LA. 252 pp. Romano III, F.A., B. Barreras-Borrero, and D.R. Nelson, 2001. Ecological distribution and community analysis of Tardigrada from Choccolocco Creek, Alabama. Zoologischer Anzeiger 240:535–541. 98 Southeastern Naturalist Vol.7, No. 1 Séméria. Y. 1982. Recherches sur la faune urbaine et semi-urbaine des tardigrades muscicoles et lichénicoles. II. L’éspace sub-urbain: Les hauteurs orientales de Nice-Ville. Bulletin de la Société Linnéenne de Lyon 51:315:328. Simpson, G.G. 1964. Species densities of North American mammals. Systematic Zoology 12:57–73. Webb, S.D. 1990. Historical biogeography. Pp. 70–100, In R.L. Myers and J.J. Ewel (Eds.). Ecosystems of Florida. University of Central Florida Press, Orando, FL. 2008 H.A. Meyer 99 Appendix 1. Distribution and morphology of tardigrades found in the state of Florida. For site information see Figure 1 and Table 1. All measurements are in micrometers. Measurements in parentheses are means (absent if only one specimen was measured). Abbreviations: L = body length, BTL = buccal tube length, BTW = buccal tube inside diameter, MPL1–3 = length of macroplacoids 1–3, mpl = length of microplacoid, I–III = length of outer claws I–III, IV = length of claw IV, OD = diameter of egg including egg processes, ID = diameter of egg excluding egg processes, PH = height of egg process, C = number of processes around egg circumference. Class Heterotardigrada Order Echiniscoidea Family Echiniscidae Echiniscus cavagnaroi Schuster and Grigarick, 1966 11 specimens. Sites: 50, 53a, 54a, 68bc. Morphology: L 108–200 (158.7), Cirrus A 16–30 (23), Spine C 3–7.5 (4.8), Spine Dd 5–16 (9), Spine E 3–12 (6.6). Echiniscus kofordi Schuster and Grigarick, 1966 6 specimens. Sites: 28a, 55d, 71, 74. Morphology: L 91–163 (123.9). Echiniscus virginicus Riggin, 1962 13 specimens. Sites: 10b, 11ab, 22c. Morphology: L 122–190 (158), Cirrus A 20–28 (24), Spine C 11–20 (15.5), Cd 2–3 (2.5), D 13–20 (15.9), Dd 3–15 (8.3), E 12.8–22.0 (17.9). No E. virginicus, including those in the adult size range, had a Spine E. This differs from the organisms studied by Christenberry and Mason (1979) elsewhere in the South, where adults, but not subadults, possessed Spine B. Pseudechiniscus suillus Ehrenberg, 1853 30 specimens. Sites: 9a, 24a, 26a, 48b, 82a. Morphology: L 116–180 (147), Cirrus A 15–21 (19). Class Eutardigrada Order Parachela Family Hypsibiidae Diphascon (Diphascon) pingue (Marcus, 1936) 2 specimens. Ramazzottius baumanni (Ramazzotti, 1962) 8 specimens. Sites: 10a. Morphology: L 260–270 (265). Family Macrobiotidae Macrobiotus echinogenitus Richters, 1903 106 specimens, 2 eggs. Sites: 8a, 11a, 18ab, 25ab, 26ab, 32a, 35, 37, 42ab, 48a, 51, 52ab, 61ab, 62, 82b. Morphology: L 138–500 (281.6), BTL 23–42 (30.9), BTW 3–7 (4.5), MPL1 4–11 (7.3), MPL2 2.8–7.5 (4.5), mpl 1–3 (1.9), I–III 5–13 (8), IV 6–13 (9), OD 83, ID 60, PH 10, C 15. Macrobiotus cf. harmsworthi sp. 1 56 specimens, 4 eggs. Sites: 14, 19, 49, 50, 65. Morphology: L 160–460 (322.2), BTL 28–40 (32.5), BTW 3.5–7 (5.5), MPL1 3–7 (4.8), MPL2 2–5 (3.9), MPL3 2.5–6 (4.1), mpl 1.5–4 (2.9), I–III 6–10 (8), IV 6–11 (9), OD 75–81 (79), ID 52–61 (56.7), PH 7–15 (10.7), C 15–24 (18). Macrobiotus cf. harmsworthi sp. 2 6 specimens. Sites: 27a. Sites: 27a. Morphology: L 260, BTL 35–40 (38), BTW 7.5– 10.5 (9.3), MPL1 4–6.5 (5.5), MPL2 4–5 (4.7), MPL3 4.8–5.5 (5.2), mpl 4–5 (4.7). 100 Southeastern Naturalist Vol.7, No. 1 Macrobiotus cf. hibiscus 7 specimens. Sites: 28a, 38b, 68a. Sites: 28a, 38b, 68a. Morphology: L 260–385 (297.8), 27–35 (30.8), BTL 3.8–5.8 (4.8), MPL1 5–8 (7), MPL2 5–5.5 (5.2), mpl 2–3 (2.3). Macrobiotus hufelandi Schultze, 1833 32 specimens, 2 eggs. Sites: 16abc, 21, 64, 74. Morphology: L 240–260 (250), BTL 28, BTW 3, MPL1 6, MPL2 4, mpl 1, OD 71–83 (77), PH 5 (5), C 26 (26). Macrobiotus cf. hufelandi sp. 1 38 specimens. Sites: 14, 19, 49, 50, 65a. Morphology: L 155–415 (335.5), BTL 26–40 (32.3), BTW 4–6.8 (5.3), MPL1 4.5–11 (8.1), MPL2 3–6 (5.1), mpl 1–3 (2), I–III 8–12 (10), IV 10–14.5 (12.4). Macrobiotus cf. hufelandi sp. 2 260 specimens, 22 eggs. Sites: 1, 5, 7b, 12ab, 15, 17, 22a, 23, 28b, 31, 40ab, 54abc, 55abc, 56a, 60abc, 67ab, 69ab, 71, 82a. Morphology: L 160–602 (335.9), 28–46 (34.9), BTW 3–7.5 (5.1), MPL1 3.5–12.5 (8.8), MPL2 3.5–9 (5.7), mpl 1–5 (2.6), I–III 6–11 (9), IV 5–12 (10), OD 68–96 (78.7), ID 56–85 (68.7), 4–6 (5), C 22–36 (27.6). Macrobiotus islandicus Richters, 1904 3 specimens. Sites: 53c. : L 255–300 (271.7), BTL 22–40 (31.3), BTW 2–6 (4.2), MPL1 5–8 (6.7), MPL2 2.5–5 (4.2). Macrobiotus occidentalis Murray, 1910 12 specimens. Sites: 8b, 58. Morphology: L 256–395 (307.1), BTL 28–36 (31.2), BTW 3.8–5 (4.4), MPL1 6–8 (7.2), MPL2 4–5 (4.7), mpl 0.5–2 (1.8), I–III 6–11 (9), IV 10–11 (10). Macrobiotus richtersi Murray, 1911 39 specimens. Sites: 16abc, 24b, 25a, 59, 63d, 77, 79, 81a. Morphology: L 355–630 (425.5), BTL 31–52 (42.5), BTW 8–11 (10.1), MPL1 5–11 (7.5), MPL2 3–8 (5.8), MPL3 (8), mpl 3 (3). Macrobiotus tonollii Ramazzotti, 1956 33 specimens, 2 eggs. Sites: 13ab, 48c, 55a, 69b. Morphology: L 420–680 (494.4), BTL 31–55 (41.9), BTW 5–11 (7.9), MPL1 4–10, MPL2 3–6 (4.8), MPL 3 5–11.5 (7.8), I–III 8–10 (9), IV 10–12 (11), OD 105–116 (109.5), ID 60–70 (55), C 8 (8). Minibiotus fallax Pilato, Claxton, and Binda, 1989 98 specimens, 3 eggs. Sites: 3, 7a, 27b, 29b, 33ab, 43ab, 44, 46, 68abc. Sites: 3, 7a, 27b, 29b, 33ab, 43ab, 44, 46, 68abc. Morphology: L 160–390 (278), BTL 21–30 (25), BTW 2–3 (2.8), MPL1 5–6 (5.5), MPL2 3–4 (3.5), I–III 9–10 (10), IV 10–11 (10). Minibiotus intermedius Plate, 1889 128 specimens, 7 eggs. Sites: 1, 2, 6, 9abc, 10ab, 11ab, 12ab, 15, 19, 22a, 25a, 26bc, 27a, 28a, 29a, 31, 32b, 34, 38ab, 48cb, 52b, 53abc, 54a, 55b, 55d, 56ab, 58, 61b, 65c, 71, 79, 81a, 82a. Morphology: L 88–232 (176.6), BTL 18–28 (20.6), BTW 1–2 (1.4), MPL1 1–2 (1.6), MPL2 1–2 (1.5), MPL3 1–2 (1.5), I–III 3–7 (5), IV 3–8 (6), OD 43–52 (47.2), ID 35–45 (40.3), PH 3–5 (4), C 30–34 (32). Order Apochela Family Milnesiidae Milnesium tardigradum Doyère, 1840 573 specimens. Sites: 1, 2, 4, 6, 7b, 8b, 13ab, 14, 16a, 17, 18b, 19, 20, 21, 22c, 25b, 27ab, 28ab, 29b, 29c, 30, 33a, 34, 35, 36, 38b, 39, 41, 43ab, 44, 45, 46, 47, 53a, 54b, 55bcd, 56a, 59, 60a, 61ac, 63abcde, 65abcd, 67b, 70, 71, 72ab, 73, 74, 76ab, 77, 78, 80, 81b, 82a, 83. Morphology: L 105-747 (443), BTL (10-70), BTW 5-21 (11.7).